JOURNAL oF te ARNOLD ARBORETUM HARVARD UNIVERSITY VOLUME 65 NUMBER 1 US ISSN 0004-2625 Journal of the Arnold Arboretum Published quarterly in January, April, July, and October by the Arnold Arboretum, Harvard University. Subscription price $50.00 per year, plus $5.00 postage for addresses outside of U.S.A Subscriptions and remittances should be sent to Journal of the Arnold Arboretum, P. O. Box 368, Lawrence, Kansas 66044, U.S. A. Claims will not be accepted after six months from the date of issue. POSTMASTER: send address changes to Journal of the Arnold Arboretum, P. O. Box 368, Lawrence, Kansas 66044, U.S. A. Volumes 1-51, reprinted, and some back numbers of volumes 52—56 are available from the Kraus Reprint Corporation, Route 100, Millwood, New York 10546, U.S.A EDITORIAL COMMITTEE S. A. Spongberg, Editor E. B. Schmidt, Managing Editor P. S. Ashton K. S. Bawa P. F. Stevens C. E. Wood, Jr. Printed at Allen Press, Inc., Lawrence, Kansas COVER: The stylized design appearing on the Journal and the offprints was drawn by Karen Stoutsenberger. Second-class postage paid at Boston, Massachusetts, and additional offices. JOURNAL OF THE ARNOLD ARBORETUM VOLUME 65 JANUARY 1984 NUMBER 1 TEMPLE CLAYTON, CHEMIST AND AMATEUR BOTANIST, 1914-1978 BERNICE G. SCHUBERT THE FIELDS of natural history and botany have always had among their ranks rather large numbers of interested and enthusiastic amateurs. In botany, at least, these people have devoted themselves whole-heartedly and often in a most professional manner to a pursuit that was actually a hobby, not the effort from which they derived their livelihood. In the past and even now, many amateurs have been and are much concerned with monocotyledonous plants, especially members of such families as the Bromeliaceae, Liliaceae, and Or- chidaceae—all taxonomically and horticulturally difficult families with species scattered in often remote, hard-to-reach, tropical areas. Temple Clayton was one of those dedicated amateurs and had a emer interest in the Dios- coreaceae, or family of the true (tropical) yam Born in Andover, New Hampshire, on ae 19, 1914, Temple Clayton died at White Bear Lake, Minnesota, on November 10, 1978. His early school- ing (1927-1931) was at the Tilton Preparatory School, Tilton, New Hampshire. From there he went to Cornell University, where he majored in chemistry and received the bachelor of chemistry degree in 1935. He then went to the New York City area, where, while working as a chemist for various companies’ and teaching chemistry,? he also did graduate work at Columbia University, re- ceiving the M.A. degree in 1941. From June, 1944, to August, 1954, Mr Clayton was employed as a group leader by the Schering Corporation, a phar- maceutical firm in New Jersey with interests in natural products. Through his work involving chemical constituents of the underground parts of various species of Dioscorea, he became interested in the plant sources. (He had earlier ‘September, 1935-June, 1936, as research chemist for Kessler Chemical Corp.; August, 1936- February, 1937, as chemist for Carleton Ellis. 2First (February, 1937-June, 1938) at Newark College of Engineering, then (September, 1938—June. 1944) at Cooper Union © President and Fellows of Harvard College, 1984. Journal of the Arnold Arboretum 65: 1-4. January, 1984. a JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 - Shy r.’. TEMPLE CLAYTON 1914-1978 become interested in plants through the influence of his uncle, the late A. Le Roy Andrews, professor of Germanic philology at Cornell, but well known for his interest in plants and as an American authority on mosses, particularly the genus Sphagnum.) As an intellectual adjunct to his chemical work, Temple Clayton plunged from a rather casual concern with local flora to a serious study of a botanically difficult plant family. He pursued this activity mostly on his own time, but with travel support from the Schering Corporation, botanical support from the New York Botanical Garden, and some technical assistance and serious interest from the U. S. Department of Agriculture. In November, 1954, Temple Clayton went to work for the 3M (Minnesota Mining and Manufacturing) Company, where from the time he arrived until July, 1962, he worked in the Central Research Laboratories, first as a literature chemist then as a patent liaison. From August, 1962, until his death in 1978, — 1984] SCHUBERT, TEMPLE CLAYTON, 1914-1978 3 Specimens of Dioscoreaceae From the Collections of Temple Clayton Presented to the Arnold Arboretum of Harvard University by the Herbarium, Department of Botany, University of Minnesota, St. Paul, 1979 Photographs of Dioscoreaceae From the Collections of Temple Clayton Presented to the Arnold Arboretum of Harvard University by the Herbarium, Department of Botany, University of Minnesota, St. Paul, 1979 Above: label to be used on Temple Clayton’s herbarium specimens. Below: label to be used on his photographs. he was employed as a patent agent in the Office of Patent Counsel of 3M. Mr. Clayton was admitted to practice before the United States Patent and Trade- mark Office, and he was a member of the Minnesota Patent and Trademark Law Association. During his 24 years in Minnesota, Mr. Clayton continued his interest in the Dioscoreaceae, supported by the encouragement of Gerald Ownbey, professor of botany and curator of the herbarium at the University of Minnesota, St. Paul. So far as I can tell, it was during this period that Temple Clayton for- mulated most of his ideas about the reorganization of the family Dioscorea- ceae—work for the most part written up, sometimes roughly and sometimes in more finished form, but none of it published. He had, however, annotated specimens in many herbaria with unpublished names. He photographed every specimen he had on loan from European herbaria, proposed a whole new classification for the family Dioscoreaceae and its com- ponents, wrote detailed descriptions for a large number of new taxa at varlous levels, and made sketches illustrating significant characters of the plants. He accumulated a vast bibliography, a good study collection of specimens, and a knowledge of theoretical approaches to taxonomic matters that was, to say the least, surprising. At Temple Clayton’s death, all the materials were given to the herbarium of the Department of Botany, University of Minnesota, St. Paul, by his wife, Emma G. Clayton. However, Professor Ownbey thought that the facilities and area of specialization there were not the best for materials concerned with an 3From a letter of Mr. Donald C. Gipple, senior patent attorney, 3M Co., March 1, 1982. 4 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 important group of tropical plants, and they were transferred to the Arnold Arboretum of Harvard University in 1979. Organizing the specimens and photographs to make them easily available and most useful has been a rather lengthy process, due to lack of continuous time, not lack of interest. The specimens (some mounted and some unmounted; some determined, some named as new, and some sterile and unidentifiable or represented by cultivated material grown by the U.S. Department of Agriculture) have been inventoried. Before being mounted, each specimen and photograph will have a special label afhxed indicating the source and date of the gift. Duplicates will be made available to institutions whose collections were studied, as well as those with special interest in the Dioscoreaceae. The acquisition of the Temple Clayton dioscoreaceous reliquiae has proven to be an interesting and somewhat awesome experience. The items making up the collection comprise the results of about 30 years of devoted attention to a fascinating avocation for which Mr. Clayton was basically untrained. 1 make this last remark not in a sense of judgment, which is not my purpose here, but in a sense of wonder that he accomplished such a large amount of time- consuming work and made so many meticulous observations in spare hours. Although the materials were turned over with “no strings attached,” Temple Clayton’s friends and colleagues in Minnesota hoped that something publish- able might be found that could bear his name as author. There is little doubt that some of the species he indicated as new, and for which he wrote descriptions and annotated specimens, were indeed undescribed. To be as certain as possible, I enlisted the collaboration of Dr. Franklin Ayala, director of the Herbarium Amazonense, Iquitos, Peru, and also a student of Dioscoreaceae, to study some of the material with me. In the short time we had to work together, we chose four species from among those left by Temple Clayton: these will be published in a separate paper. My acquaintance with Temple Clayton extended over a long period, starting before he left New Jersey for Minnesota, when he was studying plant material at the New York Botanical Garden and visiting the (then-called) Plant Industry Station of the U.S.D.A. at Beltsville, Maryland, to see some of his own material growing in the greenhouses. He was generous about sharing with me materials he had on loan from Europe, as well as information as he acquired it on Dioscoreaceae. He was an interesting, enthusiastic person, very serious about his chief interests, with an amiable personality and boundless energy. I trust that my colleagues and | will be able to do justice to some of the results he left unpublished. 4 For the photograph and other useful information I am indebted to Mrs. Emma G. Clayton; for other data, to Mr. Donald C. Gipple, of the 3M Com- pany; and for the privilege of having the material presented to the Arnold Arboretum of Harvard University for my study, to Dr. Gerald B. Ownbey. ARNOLD ARBORETUM OF HARVARD UNIVERSITY 22 Divinity AVENUE CAMBRIDGE, MASSACHUSETTS 02138 1984] ROGERS, ZINGIBERALES :) THE ZINGIBERALES (CANNACEAE, MARANTACEAE, AND ZINGIBERACEAE) IN THE SOUTHEASTERN UNITED STATES! GEORGE K. ROGERS THE ORDER ZINGIBERALES (Scitamineae, Scitaminales) is clearly delimited and is universally accepted as a natural group of four to eight families, de- pending on the ranks assigned to some of its chief components. Emphasizing anatomical evidence in addition to morphology, Tomlinson (1962, 1969) ad- vocated recognition of Zingiberaceae, Costaceae (often included in Zingiber- aceae), Marantaceae, Cannaceae, Musaceae, Lowiaceae, Heliconiaceae, and Strelitziaceae (the last three often included in Musaceae). Other recent authors differ as to which of the segregates are chosen for elevation to the rank of family. Zingiberaceae, Cannaceae, and Marantaceae are represented in the southeastern United States by a small number of aoa and introduced species. Transformation of stamens into sterile staminodes is a well-known trend bearing on the interrelationships of the families of Zingiberales. The Musaceae 'Prepared for the Generic Flora of the Southeastern United States, a long-term project of the Arnold Arboretum pie. made possible through the support of the gee Science Foundation under grants BSR-8111520 to Harvard University (Carroll E. Wood, Jr., Arnold Arboretum, principal investigator) al BSR-8303100 to the University of the State of New ey (Norton G. Miller, New York Biological Survey, principal investigator). This treatment, prepared under BSR-8111520. 1s the 101st in the series. It follows the format established in the first paper (Jour. Arnold Arb. 29: 296- 346. 1958) and continued to the present. The area covered by the Generic Flora includes North and descriptions are based primarily on the plants of this area, with information about extraregional ers of a family or genus in brackets [ ]. References that I have not verified are marked with an aster Scccal thanks are due to those who responded to my requests for information on Zingiberales. All of the replies were thoughtful and truly helpful. | am indebted in this connection to Daniel F Austin, the late George N. Avery, Carl W. Campbell, Robert K. Godfrey. Richard A. Howard, Walter S. Ju Julia F. Morton, Ghillean T. Prance, and John W. Thieret, as well as to the additional authorities consulted by some of them. Figure | was drawn by Margaret van Montfrans using dis- sachusetts. Karen Stoutsenberger, supervised by Kenneth Robertson and Carroll Wood, prepared Figure 2 from a living specimen collected in Florida by Carroll Wood and cultivated at the Arnold Arboretum in Jamaica Plain. Figures 3 and 4 are also the work of Karen Stoutsenberger, directed by Carroll Wood. Materials for these figures were collected or otherwise provided by Manuel Flores, Robert Godfrey, Richard Howard, Peter Stevens, and Carroll Wood. Stephen Spongberg and Carroll Wood are thanked for their careful reviews of the manuscript. Elizabeth Schmidt's editorial im- ments are gratefully acknowledged. Barbara Nimblett substantially expedited preparation of . einecont entries President and Fellows of Harvard College, 1984. Journal of the Arnold Arboretum 65: 5-55. January, 1984. 6 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 and its segregates have five or rarely six fertile stamens, a condition that invites interpretation as primitive (see pertinent discussion in Dahlgren). Zingiberaceae sensu lato, Marantaceae, and Cannaceae have only one fertile stamen, with the others developing as petaloid staminodes or sometimes absent; in the two latter families half of the fertile stamen is petallike. Comparison of the families is provided in the key below, where further ties between the Cannaceae and Marantaceae—and characters separating these families along with Zingibera- ceae from Musaceae sensu /ato—are enumerated. Inviting as it may be to extrapolate a phylogenetic scheme from the characters in the key, the cladistic relationships of the eight putative families of the Zingiberales are obscured by contradictory characters, uncertainties surrounding homologies of apparent similarities (e.g., the diverse “arils’” and chalazal inclusions in the seeds), and insufficient sampling for several characters. Phylogeny within the order and problems with its assessment were discussed by Tomlinson (1962), who sug- gested Strelitziaceae as the least specialized family The best taxonomic position for the Fieibeales is debatable. Hutchinson derived them from the Bromeliales (one family), and these from Commelinales. Bromeliales and Zingiberales were thought by Takhtajan to have a common origin from llialean stock: Cronquist (1978, 1981) united the two orders as subclass Zingiberidae. In Thorne’s scheme, the Zingiberales were placed in superorder Commeliniflorae along with the Bromeliaceae (a member of the Commelinales). Superorder Zingiberiflorae (containing only the Zingiberales) was hypothesized by Dahlgren (p. 140) to have arisen from pee ancestors on a “pro-commeliniflorean-zingiberiflorean-bromeliiflorean brane A prevalent concern in the literature on Zingiberales is inter ne of their androecia in terms of a presumably ancestral arrangement with three stamens in an antesepalous outer cycle alternating with three others in an antepetalous inner cycle. Interpretations involving patterns of venation, and positional. teratological, developmental, and anatomical evidence bearing on this matter are reviewed in Costerus (1916a, 1916b, 1916c), Eichler (1873: 1884. under Marantaceae), Holttum (1950, under Zingiberaceae), Kirchoff (1983b), Rao ef al., and Schachner. A brief overview follows. In the relatively uncontroversial Marantaceae the inner cycle of the androe- cium probably includes the fertile stamen with its lateral appendage, the cu- cullate staminode, and the callose staminode (synonymy for these floral parts is provided with the family description). The outer cycle must then be repre- sented by one (usually) or two petallike staminodes, with the third member suppressed. Support for this widely accepted interpretation appears in devel- opmental studies by Eichler (1884) and Kirchoff (1983b): both observed a primordial ring, positioned above the calyx, which splits tangentially in each of the three alternisepalous positions to yield altogether three petals to the outside and three members of the androecium (fertile stamen, cucullate stam- inode, and callose staminode) to the inside. Primordia for the remaining (outer) staminodes are antesepalous. Eichler found floral vasculature to corroborate the developmental evidence (see also Tilak & Pai, 1966, 1968). On the basis of developmental similarities, studies of Eichler (1884) and Kirchoff (1983b) agree further that flowers of Cannaceae are fundamentally as — enon —_— 1984] ROGERS, ZINGIBERALES f, described above for Marantaceae, with one or two (or sometimes no) petallike staminodes representing the outer androecial cycle, and with the labellum, one petallike staminode, and the fertile stamen (or no) as the inner cycle. According to both authors, these structures are homologous with the marantaceous cu- cullate staminode, callose staminode, and fertile stamen, respectively. Mar- antaceae and Cannaceae have been said to differ in the sequence of initiation of calyx lobes: those of Cannaceae spiraled in the same direction as the corolla and androecium vs. the spirals opposed in Marantaceae. However, Costerus (1916c) observed in a species of Canna many instances in which the calyx and corolla formed opposite spirals. Few contemporary observers would dispute that in the Zingiberaceae the inner staminal cycle is represented by two staminodes connate to form the labellum plus the separate fertile stamen; clearly the outer cycle is sometimes (e.g., in Hedychieae) manifest as two free petaloid staminodes. However, the fate of the third (median) outer staminode, which is never unambiguously apparent, and the whereabouts of the two lateral staminodes when they are not obvious, remain problematic. Authors commonly consider fusions with the labellum as potentially accounting especially for the recondite median staminode (e.g., see Costerus, 1916b; Gregory), and sometimes for all three outer staminodes. Pai (1965b, 1966), however, stressed vascular evidence, while arguing in favor of the “classical view” that the labellum in at least some Zingiberaceae involves only the two inner staminodes. At the other extreme, the labellum composed of all five staminodes has been numbered among the characters that distinguish Costus L. (-oideae, -aceae) (see Costerus, 1916b; Cronquist, 1981; Holttum, 1950; Loesener; Maas, 1972; Schachner; Thomp- son, a contrary conclusion; Tomlinson, 1956: several of the references in this and the following paragraph are listed under Zingiberaceae). This distinction is best regarded as open to refinement. In some genera of Zingiberoideae, the lateral staminodes are ostensibly manifest as large or small lobes or teeth on the labellum. Dampening the temptation to regard all lobes on the labellum in Zingiber- aceae as adnate outer staminodes, Burtt (1972) pointed out that the elaborately lobed labellum of some orchids is homologous with a single perianth member. He also observed that the number of lobes on the zingiberaceous labellum in some cases exceeds the number of missing staminal components. Moreover, in Hedychium coronarium, in which the lateral staminodes are strongly and distinctly developed, the labellum sometimes has lateral lobes. ontrary to interpretations of the epigynous glands of Zingiberaceae as mod- ified stamens or styles, these nectaries have been shown to be vascularized carpellary outgrowths that range in position from being enclosed within apically open ovarian cavities in Costoideae to rising vertically above the summit of the ovary in Zingiberoideae (Pai, 1966, Rao). GENERAL DESCRIPTION OF ZINGIBERALES: Small to arborescent, perennial, rhizomatous, mostly terrestrial herbs typi- cally of moist tropical habitats. Hairs mostly unicellular. Leaves usually pet- 8 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 iolate, with sheathing bases (these often forming a pseudostem around a feeble true stem); blade entire, rolled in the bud, with a multistranded midrib region and many pinnate-parallel, lateral nerves; leaf axis usually with septate air canals. Inflorescences often with large, folded to spathelike, colorful primary bracts. Flowers usually perfect, zygomorphic or asymmetric, typically showy. Sepals 3, unlike the petals; petals 3. often unequal. Androecium the showiest portion of the flower (in our families), arranged fundamentally in 2 trimerous cycles; fertile stamen | [or 5 or 6], when | the flower with showy staminodes, these diverse in number, form, arrangement, and fusions, usually showier than the perianth. Gynoecium 3-carpellate, syncarpous and basically 3-locular (2 locules sometimes aborting), with | to many ovules per locule, ovary inferior [or flowers perigynous], style single. Seed with a specialized region of dehis- cence, often operculate, usually with a micropylar collar, usually arillate (or with arillike emergences from the funiculus). KeEY TO THE FAMILIES AND GENERA OF eG RALES IN THE SOUTHEASTERN UNITED STATES A. pees ee 5 [or 6]; raphide-sacs present: guard cells with the inner and er ledges equal. Musacrae (plants often of treelike appearance and size, with ie buds see the leaves in Musa). 000 ee [Musa.3} netional stamen 1; raphide-sacs absent; guard cells with the inner and outer ledges unequal (except in Cannaceae B. Leaf sheaths ligulate ease 1. b); flowers zygomorphic; sepals connate into a tube usually slit along | side (FiGure 1, g); sterile staminodes represented by a usually broad peti um composed of 2 or more fused staminodes (the dual nature not readily evident) flanked by and sometimes adnate to 2 petallike to much reduced (to not detectable) es staminodes; anther with 2 locules, the style lying between them (FiGure 1, h), ona emoupe! taloid named pollen not deposited onto style in bud: nectaries vasculari emergences of carpellary tissue [or sometimes concealed in cavities]: ae development helobial. Zinci- BERACEAR. Lateral staminodes strongly developed and petallike (FiGURE 1, g); labellum more or less flat (not troughlike), white to yellowish or eee with yellow: corolla tube several centimeters long; none of petals ome pedo teat Seay aang ec gue cehapeecatig dun Gita act enue) we ates ded aires eae cin teats Hedychium. Lateral staminodes not apparent or reduced to small teeth: ee trough- like, yellowish (especially toward the margin) or white, with reddish markings (especially toward hg center): corolla tube to ca. | cm long; | petal hoodlike and facing labelam, .. ec s void eedqdiuchin ne hahah ane veea aces es [Alpinia.*] Leaf sheaths without Tieulec (sometimes auriculate); flowers asymmetric; sepals distinct; sterile staminodes variously arranged but all distinct (except for basal (=) a *For embryological comparisons of these families, see Panchaksharappa (under Zingiberaceac). ‘Species of Musa L. are sometimes found growing outside of cultivation in Florida (and, according to Small, in Louisiana). Such occurrences appear to en usually, iP nol fee from persistence after cullivation, from rhizomatous spreading, or {rom 1 vertent di n by humans. Available evidence does not confirm bananas as truly naturalized panes ey of the flora of the southeastern United States (see ae & Black, Lakela & Craighead, Long & Lakela, Melvill, Poppleton er al., Thieret, Ward, Wunderlin, and acknowledgments in footnote 1). “For comments on A/pinia Roxb., nom. cons., in the southeastern United States, see Zingiberaceae. 1984] ROGERS, ZINGIBERALES 9 fusions); anther with | locule, the remainder of stamen petaloid or partly so; pollen aa onto style in bud; nectaries septal; endosperm development nuclea D. Petioles not pulvinate at apex; stem with mucilage canals; staminodes con- sisting of a labellum and [0 or] 2 or 3 petallike staminodes (none of these hoodlike or appendaged) (FiGure 2, d, e); style more or less petaloid, not curling inward, the region of pollen deposition not specialized, the stigma apical and marginal (FiGure 2, k); gynoecium 3-locular, with many ovules per locule; seed opening by a slit, without canals in the (mostly chalazal) nutritive tissue; embryo straight; flowers single or in homodromous pairs. CANNAGEAB ods UG Sore ha nantes Ma ee ge ae 1. Canna. D. Petioles pulvinate at apex: stem without mucilage canals; staminodes con- sisting of an appendaged, pouchlike “cucullate” staminode (FiGure 3, g) enclosing the style before ala en. a hoodlike “‘callose” staminode (FIGURE 3, h), and 1 or 2 petallike staminodes; style not petaloid, curling rapidly inward when dislodged and effecting pollination, with a specialized region of pollen deposition (FiGure 3, k-13), the stigma in a cleft (FiGure 3, k-14); gynoecium unilocular (in ours), with 1 ovule per locule; seed operculate, with 1 or 2 canals in the (nucellar) nutritive tissue; embryo bent; flowers in mirror- image Sais pairs (this not always conspicuous in Maranta). Ma- RANTACEA E. Petals eee or nearly so; cucullate staminode with 2 appendages (FIGURE 3, g); outer petaloid staminode | (FiGure 3, f); gach cleft with a bifid rim projecting back into the flower (FiGure 3, i-k); axes of flower pairs condensed (flowers of a pair borne tightly side by side; ee RE 3, b); canal in seed forked at the base (in effect 2 canals). .............. l. Thalia. Petals connate into a tube; cucullate staminode with | appendage; outer petaloid staminodes 2: stigmatic cleft more or less funnel shaped; axes of flower pairs elongate; canal in seed forked only at the apex ~ m™ REFERENCES: BENTHAM, G., & J. D. Hooker. Scitamineae. Gen. Pl. 3: 636-657. 1883. [Treatment of Marantaceae severely criticized by SCHUMANN (see Marantaceae). Bisson, S., S. Gutttemer, & J. L. HAmet. Contribution a étude caryo-taxinomique des Scitaminées. (English summary.) Mém. Mus. Hist. Nat. Paris, IIB. 18: 59-145. pls. 9-19. 1968. [Includes tables of chromosome numbers from other literature; . = 9 in Seas not upheld by Maas, Fl. Neotrop. Monogr. 18: 1-218. 1977. Back, D. _& §. Brack. Plants of Big Cypress National Preserve: a preliminary a of vascular plants. S. Florida Res. Center Rep. T-587. 28 pp. Homestead, Florida. 1980. [Musa = paradisiaca, 7; uncommon. Corre.t. D. S., & H. B. Corrett. Aquatic and wetland plants of southwestern United States. xv + 1777 pp. Environmental Protection Agency, Washington, D. C. 1972. [Reissued in 2 volumes by Stanford Univ. Press. 1975; Canna indica, C. flaccida, C. a Thalia dealbata, 684-687.] CosTerus, J. A fresh Saree into the structure of the flower of Canna. Ann. ard. Bot. Sen Il. 14 -184. pls. 27, 28. 1916a. [Includes diagrams, dis- sections, and anatomical ae of flowers of Canna; describes 2 rudimentary styles (dubious) and teratologies.] Das Labellum und das Diagram der Zingiberaceen. /bid. 14: 95-108. pl. 17. 1916b. [Teratological and anatomical evidence that the labellum involves at least 2 inner staminodes plus tissue from the outer whorl; see comments on this paper in SCHACHNER; Cf. Pat, 1965b — a 10 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 —. Die Ubereinstimmung und der Unterschied in dem Bau der Blumen von Canna und derjenigen der Marantaceen. [hid. 15: 59-93. pis. 13, 14. 1916c. [Includes a point-by-point comparison of the families; favors the unusual view that the partly petaloid stamen ‘so wohl bei Marantaceen als bel Canna aus zwei genetisch wn- abhdngigen, aber spaiter verwachsenen Teilen (1 Stamen und | Staminodium) be- steht” (p. CRONQUIST, A. The Zingiberidae, a new subclass of Liliopsida (monocotyledons). Brit- tonia 30: 505. 1978. [Comprises orders Zingiberales and Bromeliales, elaboration in the following reference. ] . An integrated system of eee of flowering plants. xvii + 1262 pp. New York. 1981. [Zingiberales, 1165-1188: includes comparison of Zingiberidae with Commelinidae and Liliidae. and comparison of Bromeliales with Zingiberales (pp. 1158, 1159). Cutten, J. William Roscoe’s monandrian plants of the order Scitamineae. Notes Bot. Gard. Edinburgh 32: 417-421. 1973. [Includes biographic information on Roscoe, eae: of publication of the work (issued in 15 parts, 1824-1829), and index to — plates. ] Czaja, A. T. Structure of starch grains and the classification of vascular . lant families. Taxon 27: 463-470. 1978. [Lists 14 types of starch grains; seeds of Cannaceae, Marantaceae, Zingiberaceae. and other. monocots store starch primarily in highly compound starch grains. DAGHLIAN, C. P. Zingiberalean leaves from the Lower Eocene of Texas. (Abstract.) Bot. Soc. Am. Misc. Publ. 162: 57, 58. 1982. [Probably related to Cannaceae, cf. Hickry & Pererson and FriepricH & Kocu (1970, 1972) (all under Zingiberaceae).] DaHLGreN, R. General aspects of angiosperm evolution and macrosystematics. Nordic Jour. Bot. 3: 119-149. 1983. [Includes discussion of placement of Zingiberiflorae (Zingiberales).] Davis, G. L. Systematic embryology of the angiosperms. vill + 528 pp. ” York, London, & Sydney. 1966. [Cannaceae, 73; Marantaceae, 170, 171; Snes 277, 278. DuNCcAN, WwW. H., & J. T. KArresz. Vascular flora of so hae An annotated checklist. ix + 146 pp. Athens, Georgia. 1981. [Canna flaccida, C. x generalis, C. indica, Thalia dealbata, 44, Hedychium not listed. ] Eicuier, A. W. Uber den Bliithenbau ve Canna. Bot. Zeit. 31: 177-189, 193-198, —218, 225-232, 241-247. p/. 2. 1873. [Includes detailed description of inflores- cence (but fig. /2 appears to be mislabeled); see Ercuier (1875; and 1884, under Marantaceae) for changes in opinion pane floral morphology. ] Scitamineae. Bliithendiagramme 1: 167-178. 1875 ErptMAN, G. Pollen morphology and vate aes (Corrected reprint of the 1952 edition, with a new addendum.) xiii + 553 pp. New York. 1971. [Cannaceae, 95, 96; Marantaceae, 264; Zingiberaceae, o 455.] GIBBS, R. D. Chemotaxonomy of flowering plants. Vol. 3. Pp. 1275-1980. Montreal & eine 1974. [Zingiberales, 1971-1980 (see also 4: 2371, 2372): current knowledge order “inadequate for serious chemotaxonomic discussion” (p. 1976): the order rich in phenolic acids (listed p. 1978); hardly surveyed for saponins: few alkaloids known (indole in Hedychium);, Zingiberaceae (includes long list of terpenoids and sesquiterpenoids; species with leucoanthocyanins listed; suggestion that Zingibera- ceae sensu stricto might stand apart in havi ng flavonoids with no B-ring substitu- tion—but see Wittiams & HARBORNE, 26, 227: flavanones, flavonols, and flavanonol listed); Senet poorly known Peat (chelidonic acid in | species. anthocyanin in | species): Marantaceae poorly known aga ceoouee by earlier author in | genus; some species possibly with leucoanthocyan ] Goperery, R. K., & J. W. Wooten. Aquatic and wetland plants of a eee United 1984] ROGERS, ZINGIBERALES 1] States. Monocotyledons. x + 712 pp. Athens, Georgia. 1979. [Canna flaccida, C. indica, C. x generalis, Thalia dealbata (illustration from Correct & CorreLt), 7. geniculata, 619-622. Greacory, P. J. The floral morphology and cytology of Elettaria cardamomum Maton. Jour. Linn. Soc. Bot. 50: 363-391. pls. J/-173. 1936. [Evidence from vasculature that labellum is composed of two inner staminodes plus one outer staminode: cf. Par (1965b), RAo et al.] HeGNAUER, R. Chemotaxonomie der Pflanzen. Band 2. Monocotyledoneae. 540 p Basel & Stuttgart. 1963. [Cannaceae, 113-116; Marantaceae, 359-361; Zinpibera. ceae, 451-471; a of Scitamineae compared chemically, 471; see also Gisss, WILLIAMS & HARBC Hepper, F. N. Cannaceae. Tn: J. HurcHinson & J. Davzie., Fl. W. Trop. Afr. ed. 2. 3: 79. 1968. [Cannas in Africa regarded as C. indica naturalized from tropical America, cf. KRANZLIN (under Cannaceae); Thalia, 85, 86. Horaninow, P. Prodromus monographiae Scitaminearum. 45 pp. 4 p/s. Leningrad. 1862. [The “Index Generum et Sectionum” includes genera and other groups of eee taxonomic ranks and thus appears not to be an acceptable designation of ein maa groups as sections in a modern sense. ] aie J. E. The development of the seed in the Scitamineae. Ann. Bot. 10: pls. 1-4. 1896. [Includes observations on Canna indica and Thalia dealbata, com- parison of families, and comparison of Costus (2 species) with other can ueeeee (3 species from 3 genera). These differ in formation of the aril and in a /O the micropylar collar and lid (but see GRooTJEN & BOUMAN (under meas and Mauritzon for indications that comparisons using these structures were pre- pane mature). | HUTCHINSON, a Zingiberales. Fam. Flowering Pl. ed. 3. Pp. 718-731. London. 1973 KiRCHOFF, B. Allometric growth of the flowers in five genera of the Marantaceae and in ae (Cannaceae). Bot. Gaz. 144: 110-118. 1983a. . Floral organogenesis in five genera of the Marantaceae and in Canna (Canna- ceae). Am. Jour. Bot. 70: 508-523. 1983b. Knutu, P. Handbook of flower pollination. English translation by J. R. AINSWORTH Davis. Vol. 3. Frontisp. + iv + 644 pp. Oxford. 1909. (Canna, Maranta, Thalia, 422, 423.] LAKELA, O., & F. C. CRAIGHEAD. Sar tae cep one vascular plants of Collier, Dade, and Monroe counties, ere . Lab. Univ. S. Florida oe 1S. viii + 95 pp. Coral Gables, Florida. 1965. ae pases around ancient dwellin Languas speciosa (Alpinia Zer eae Canna flaccida, C. indica, Thalia ae 26, 27.] Lona, R. W., & O. Laketa. A flora of tropical Florida. xvii + 962 pp. Coral Gables, Florida. 1971. [Thalia geniculata, Canna flaccida, C. indica, C. x generalis, Musa Sapientum, sas 305.] Lupsock, J. A contribution to our knowledge of seedlings. Vol. 2. 646 pp. New York. 1892. ee ce 566-569. Mananty. H. K. A cytological study of the Zingiberales with special reference to their taxonomy. Cytologia 35: 13-49. 1970. [Includes brief review of cytological literature, idiograms, drawings and photos of chromosomes, tables of chromosome numbers, and discussion of subdivision of Musaceae sensu /ato, suggests that x ultimately = 11 in Zingiberales. ] Martin, A. C. The comparative internal morphology goa Am. Midi. Nat. 36: 513- 660. 1946. (Thalia dealbata, Canna flaccida, 548, 54 MAURITZON, - Samenbau und Embryologie einiger ee een. Lunds Univ. Arssk I. Sect. 2. 31(9): 1-31. 1936. [Canna, Maranta, Calathea, Stromanthe, Hedychiwm, Ean es Kaempferia, Roscoea, Globba, Burbidgea, Alpinia, and Costus; in- JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 65 cludes details of many aspects of ovules, seeds, and embryos, especially useful in connection with the operculum and collar: embryological characters set Cannaceae apart from other Deane ] MeL VILL, J.C. List of the phanerogams of Key West. South Florida. Mem. Manchester . Philos. Soc. II]. 8: 138-154. 1884. [\fusa, 154: “sometimes found apparently ae eey | Morton, J. F. Pestiferous spread of many ornamental and fruit species in South Florida. Proc. Florida State Hort. Soc. 89: 348-353. 1976. ete speciosa, Canna indica, Hedychium coronariumn, Maranta arundinacea, 351, 352.) Atlas of medicinal plants of Middle America. xxviil + 1420 pp. Springfield, Hingis 1981. [Canna ey C. edulis, C. indica, 111,112: Maranta arundinacea, 113; Zingiberaceae, 103-107 (Costaceae, 108-1 10).] Miter, H. The fertilisation - flowers. (Translated into English and edited by D. W. THOMPSON.) xii + 669 pp. London. 1883. [Canna, Maranta arundinacea, Thalia dealbata, Hedychium, 542, 543 Neat, M. C. In gardens of Hawaii. Pies Mus. Spec. Publ. 50. x1x + 924 pp. New and rev. ed. 1965. (Third printing. 1975.) [Cannaceae, Marantaceae, Zingiberaceae, 251- is Nerouirzky, F. Anatomie der Angiospermen-Samen. Handb. Pflanzenanat. II. Arche- gon. 10. vi + 365 pp. Berlin. 1926. [Cannaceae. Marantaceae, Zingiberaceae, 87- 91.) OLaruNn, O. A. The structure and development of stomata in some Zingiberales. Notes Bot. Gard. Edinburgh 38: 499-516. 1980. [Development of stomata in all studied Zingiberales is perigenous. |} Par, R. M. Morphology of the flower in the Cannaceae. Jour. Biol. Sci. 8: 4-8. 1965a. [Discussion includes Marantaceae and Zingiberaceae ne floral anatomy of Elettaria she Maton, a re-investigation. New Phytol 64: 187-194. 1965b. [CF Grec e floral anatomy of Kaemafinie rosea Schweinf. ex Benth. with special ref- erence to the glands in Zingiberaceae. Proc. Indian Acad. Sci. 64: 83-90. 1966. & .TitAk. Septal nectaries in the Scitamineae. Jour. Biol. Sci. 8: 1-3. 1965. [Septal nectaries in Cannaceae, Marantaceae, and Musaceae, but not Zingiberaceae: although often designated as such, nectaries in Costoideae are not true septal nec- Perry, L. M. (with the assistance of JupirH MetzGer). Medicinal plants of East and Southeast Asia: attributed properties and uses. x + 620 pp. Cambridge, Massachu- setts, & London. 1980. [Cannaceae, 68; Marantaceae, 257; ee 436-444. Hedychium coronarium and other species of Hedychium, 441, Perersen, O. G. Musaceae, Zingiberaceae. Cannaceae, Marantaceae. a raat: I. 6: 1-43. 1888. [Maranteae and Phrynieae interpretable as tribes only by the - ce of the names; more explicitly designated as tribes by Schumann ao thereafter. ] fusaceae, es Cannaceae, Marantaceae. /n. C. F. P. von MARTIUS, FI. Brasil. 3(3): 1-172. a 1-50. 1890. Popeteton, J. E., A. G. Suury, & H. C. Sweet. Vegetation of central Florida’s east coast; a checklist of the eae plants. Florida Sci. 40: 362-389. 1977. [Thalia geniculata, Canna flaccida, C. x generalis, Musa saptentum, Alpinia Zerumbet, A. officinarum (p. 373). Porztat, E. Re Scitamineae. /n; H. Metcuior, A. Engler’s Syllabus der Pflanzen- familien. ed. 2: 607-613. 1964. PURSEGLOVE, J. _- ‘Tropical crops. Monocotyledons | & 2. x (+ vi) + 607 pp. in 2 vols. London. 1972. [C anges 92.93: Marantaceae, mostly aranta arundinacea 335-342; Zingiberaceac, 52 0. 1984] ROGERS, ZINGIBERALES sR: Rao, A. S., & D. M. Verma. Materials towards a monocot flora of Assam—II. (Zin- giberaceae & Marantaceae). Bull. Bot. Surv. India 14: 114-143. 1972 [1975]. Rao, V. S. The epigynous glands of Zingiberaceae. New Phytol. 62: 342-349. 1963. [Typically 2 in antero-lateral positions above the ovarian septa.] & N. Donpe. The floral anatomy of Canna flaccida. Jour. Univ. Bombay, II. 24(3)(Biol. Sci. Med. 38): 1-10. 1955. [The floral anatomy of Canna flaccida differs considerably from that of the investigated members of the Zingiberaceae”’ (p. 9).] & K. Gupte. The floral anatomy of some Scitamineae—part IV. Jour. Univ. Bombay. II. 29(3, 5)(Biol. Sci. Med. 48, 49): 134-150. 1961. [Six genera of Zingi- beraceae. ] _H. Karnik, & K. Gupte. The floral anatomy of some Scitamineae—part I. Jour. Indian Bot. Soc 33: 118-147. 1954. [Hedychium coronarium and 3 other species of Zingiberaceae; oes probably a double structure belonging to the inner whorl. ] ._M. Pat. The floral anatomy of some ee II. Jour. Univ. Bombay, II. te Sci. Med. 46): 82-114. 1959. [Four genera of Zingiberaceae: “The reduction and modification in the androecium of Zingiberaceae completely obscures the limits between the two androecial whorls” (p. 111).] HI. bid. 28(5)(Biol. Sci. Med. 47): 1-19. 1960. [Two species eee. Rickett, H. W. Wild flowers nee United States. Vol. 2. The southeastern states. Part 1. x + 322 pp. New York. 1967. [Cannaceae, Marantaceae, Zingiberaceae, 91-93; color photos of Hedychium coronarium, Canna flaccida, C. x generalis, Thalia geniculata.| Roscort, W. Monandrian plants of the order Scitamineae, chiefly drawn from living specimens. Liverpool. 1824-1829. [Descriptions and colored plates of 112 species and varieties on unnumbered pages; includes several species in the southeastern United States; see also CULLEN, and Roscoe’s preliminary paper in Trans. Linn. Soc. 8: 330-357. pl. 20. 1807. Saab, S. L., & R. K. IpRAniM. Palynological and biochemical studies of Scitamineae. Jour. Palynol. 1: 62-66. 1965. [Hedychium coronarium, Canna indica, Maranta sp.. and Strelitzia reginae, all have similar patterns of phenolic acids from the pollen (these listed) and pollen grains of similar eee (observed at low resolution); includes illustrations of sectioned grains; cf. RowLtey & SKVARLA (1974), SKVARLA & Row ey (both under Canna), and ERbTMAN & Pr: AGLOWSKI (under Hen ial ] Sato, D. The karyotype analysis in Zingiberales with special reference to the p karyotype and stable karyotype. Sci. Pap. Coll. Gen. aoe Univ. Tokyo 10: 05. 243. 1960. [Canna edulis, C. glauca, C. a VMaranta arundinacea, and extralimi- tal species: includes table of chromosome numbers from literature and table showing frequencies of base numbers arranged by genera; multiples of 9, 11, 12 common, chromosomal morphologies described and illustrated. ] SCHACHNER, J. Beitrége zur Kenntnis der Bliiten- und Samenentwicklung der Scitami- SCHERMERHORN, J. W.. & M. W. Quimsy, eds. The Lynn index, a bibliography of phytochemistry. Monograph IIL. Order, Scitamineae including the following fami- lies: Cannaceae, Marantaceae, Musaceae, Zingiberaceae; Order, Microspermae in- cluding the following family: Orchidaceae. 32 pp. Boston. SHARMA, A. Chromosome census of the plant kingdom. I, Monocotyledons. Part I. Butomales to Zingiberales. Nucleus Calcutta 15(appendix to pt. 2): 1-20. 1972. SINGH, V., & P. C. PANDE. A smudy of leaf epidermal structures in some Scitamineae. Geobios 4(1): 18-20. 1977 14 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 SkutcH, A. F. Unrolling of leaves of Musa Se Ms some related plants and their reactions to environmental ote Bot. Gaz. 90: 337-365. 1930. [Cannaceae, Mar- antaceae (Calathea), 360-36 SMALL, J. K. Manual of the eee flora. xxii + 1554 pp. New York. 1933. printed, Chapel Hill, North Carolina. 1961.) [Scitaminales, 358-362; \/usa ae tum, M. Cavendishii, Canna flaccida, C. ie a, Languas speciosa (Alpinia Ze- rumbet), Maranta arundinacea, Thalia dealbata, T. geniculata, Hedychium “coronatum.” STANDLEY, P. C., & J. A. STEYERMARK. Musaceae, Zingiberaceae. CANNACEAE, and M antaceae. ip FI. Guatemala. Fieldiana Bot. 24(3): 178-221. 195 "Hedyehium bella 198: Canna edulis, C. x generalis, C. indica, and nie species, 203- ; Maranta sil ci M. divaricata, 216-218: Thalia geniculata, 221. Teas AjAN, A. Outline of the classification of flowering plants (Magnoliophyta). Bot. Rev. 46: 225-359. 1980. [Superorder Zingiberanae (only order, Zingiberales), 318, 319: Zingiberales compared with Liliales and Commelinales Turerer, J. W. Checklist of the vascular flora of Louisiana. Part 1. Ferns and fern allies, ficken a, gymnosperms, and monocotyledons. Lafayette Nat. Hist. Mus. Tech. Bull. 2. 11 48 pp. 1972. eee C. glauca, C. indica, ‘Thalia dealbata, 37; Musa not listed. THompson, J. M. Studies in advancing sae Pt. 6. The theory of scitaminean flow- ing. Publ. Hartley Bot. Lab. Liverpool 11: 1-111. 1933. [See comments in ANDERSSON (1976, under eae. KircHuorr (1983b), and Pat (1965b). — .R.F. Proposed new realignments in the angiosperms. Nordic Jour. Bot. 3: 85- . 1983. [Zingiberales, 111; suborders Musineae and Zingiberineac.] TILAK, a D., & R. M. Pai. Studies in the floral morphology of the Marantaceae. Vascular anatomy of the flower of Schumannianthus virgatus Rolfe, a“ special reference to the labellum. Canad. Jour. Bot. 44: 1365-1370. 1966. Hl. Vascular anatomy of the flower in two species of the genus Phryvnium Willd. a Indian Acad. Sci. 68B: 240-249. 1968. III. Vascular anatomy of some species of the genus Calathea. Marathwada Univ. Jour. Sci. 2: 31-41. 1970.* TomLinson, P. B. Phylogeny of the Scitamineae— morphological and anatomical con- siderations. Evolution 16: 192-213. 1962. [This and the following paper include tables comparing systems of classification eames within this order: discusses pos- sible evolutionary trends in morphological chang . Commelinales-Zingiberales. Jn: C. R. Metrcacre, ed., evaed aes mono- cotyledons. Vol. 3. xx + 446 pp. Oxford. 1969. ite annaceae, Marantaceae, Zingi- beraceae-Costaceae, 341-389; classification of the Zingiberales nee with special reference to anatomical evidence, 295-302. | VENKATASUBBAN, K. R. A preliminary survey of chromosome numbers in Scitamineae of Bentham and Hooker. Proc. Indian Acad. Sci. 23B: 281-300. 1946. [Most of the species listed as Maranta do not belong to this genus, if their epithets are correctly applied. ] Warp, D. B. Checklist of the vascular flora of Florida. Part |. Univ. Florida Agr. Exp. Sta. Tech. Bull. 726: 1-72. 1968. [Musa acuminata (M. Cavendishii), M. paradisiaca (M. sapientum), Alpinia speciosa, Se aad edulis, C. flaccida, C. indica, Maranta arundinacea, Thalia genic ulat 56, Wiiiiams, C. A., & . Harporne. The leaf flay aneids a Zingiberales. Biochem. Syst. Ecol. 5: 221 —229. 1977. [Zingiberales very much neglected from the chemical viewpoint; flavonoid evidence shows limited similarities to Bromeliales and **Flu- viales”; includes table comparing families for leaf flavonoids: Cannaceae poor in flavonoids (based partly on second-hand reports, with cyanidin 3-rutinoside, quer- cetin, kaempferol); Marantaceae with “all classes of flavonoid anne > del- phinidin 3-rutinoside possibly common in leaves of Marantaceae but unknown in Zingiberaceae; Maranta with both flavones and flavonols: Thalia ee (from — 1984] ROGERS, ZINGIBERALES 15 other literature) with quercetin, cyanidin; Zingiberaceae with proanthocyanidins, mpferol, quercetin, myricetin, isorhamnetin, syringetin—the last 3 flavonols mostly restricted to Hedychieae; see also Gipss, HEGNAUER.] Woopson, R. E., Jr., & R. W. Scuery. Zingiberaceae, Cannaceae, Marantaceae. /n. FI. Panama. Part III. Fasc. |. Ann. Missouri Bot. Gard. 32: 57-105. 1945. [Severe criticism of Krinzlin’s revision of Canna; Canna glauca, C. flaccida, C. indica, C. edulis, 76-78, Maranta arundinacea, 101; Thalia geniculata, 104.] WUNDERLIN, R. P. Guide to the vascular plants of Central Florida. 472 pp. Tampa, Florida, & other cities. 1982. [A/usa paradisiaca (persisting after cultivation and locally spreading vegetatively), A/pinia Zerumbet (rarely See re aaa sites), Hedychium coronarium, Canna (3 spp.), Thalia geniculata, 143, | a als 67 Lindley, Key Struct. Phys. Syst. t..69..1835, nom, cons (GINGER FAMILY) Aromatic, small to large herbs with short, distichous-scaly, thickened [or long, thin] rhizomes, these branching sympodially from axils of scale leaves near the bases of erect [or sometimes very short] stems, each branch potentially terminating in an unbranched leafy shoot [branched in some Costoideae] and/ or an inflorescence. Hairs unicellular (Zingiberoideae). Plants with aromatic oil cells (Zingiberoideae). Leaves basal or cauline, sometimes petiolate, disti- chous [spirally arranged in Costoideae]; sheaths overlapping (in Zingiberoideae often forming a pseudostem surrounding or overtopping a thin true stem), open (or closed], adaxially ligulate at insertion of petiole; blade often asymmetric; stomata mostly tetracytic (paracytic to polycytic). Inflorescence(s) terminal [sometimes falsely appearing to be lateral], borne on a leafy shoot [or on a peduncle covered by distichous, bladeless sheaths arising from the rhizome], usually simple [sometimes branched or uniflorous], often with a condensed eadlike or conelike appearance, the main axes bearing spirally arranged [or rarely distichous] primary bracts subtending variably modified [sometimes 1-flowered], condensed cincinni, these with variable higher-order bract(eole)s, sometimes with basal 2-keeled prophylls. Flowers perfect [rarely imperfect], highly diverse in appearance, usually lasting only | day, mostly zygomorphic. Sepals connate into a tube (sometimes not readily distinguishable as 3 units), the tube usually split along | side and lopsided. Petals fused basally with the androecium into a usually narrow tube, the dorsal lobe usually larger than the other lobes [and sometimes appendaged]. Androecium highly modified, vari- able in appearance, composed of | fertile stamen; a usually large, petaloid, often 2-lobed labellum (usually the largest organ of the flower); and sometimes 2 petaloid [to toothlike] lateral staminodes more or less free [or fused to la- bellum]. Anther with 2 locules [sometimes variably broadened or appendaged]: pollen grains 2-nucleate when shed, usually inaperturate (Zingiberoideae), spi- nuliferous to more or less psilate. Gynoecium usually with 2 variable “‘epig- ynous glands” (nectaries) jutting into the floral tube apically from the ovary (Zingiberoideae); ovary 3-locular with axile placentation [or l-locular with parietal, basal, or free central placentation, rarely 2-locular]; ovules more or 16 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 less anatropous, usually numerous; style thin, usually passing through a channel in the filament and between the locules of the anther; stigmas diverse in form, usually ciliate, often sunken. Fruit capsular [or indehiscent, then fleshy or dry, sometimes opening by disintegration of the walls]. Seeds usually arillate, oper- culum well {to poorly] developed, endosperm development helobial, with peri- sperm and endosperm (sometimes with variable intrusions of chalazal tissue); embryo straight, cylindrical, or club shaped. Megagametophyte (embryo sac) of the Polygonum type (Zingiberoideae) [Adoxa type reported in Costus], base chromosome numbers variable, including 9, 11, 12. (Including Costaceae Nakai, Jour. Jap. Bot. 17: 203. 1941: Alpiniaceae Small, Man. Southeast. Fl. 360. 1933.) Type Genus: Zingiber Boehmer, nom. cons. A pantropical (to subtropical) herbaceous family, comprising over 1000 species in some 40-50 genera (over a third of these with very few species), typical of floors of lowland forests. Subfamily Zingiberoideae, encompassing by far the greater part of the family and subdivided into four tribes, 1s centered in tropical mainland Asia and the Malay Archipelago including Papuasia; only Renealmia L. f., nom. cons. (ca. 75 spp.), extends naturally to the New World. (It also occurs in Africa.) Subfamily Costoideae (Meisner) K. Schum. is made up of the pantropical Costus (80° spp.), the small genera Dimerocostus Kuntze and Monocostus K. Schum. in tropical America, and 7apeinochilos Miq., nom. cons., which ranges from eastern Indonesia to Queensland. No extant species of Zingiberaceae 1s indigenous to the continental United States. In Florida A/pinia Zerumbet (Pers.) B. L. Burtt & R. M. Sm. (4. speciosa (Wendl.) K. Schum.; Languas speciosa (Wendl.) J. K. Small), of tribe Alpinieae, persists after (or infrequently escapes from) cultivation, and 4. officinarum Hance reportedly persists after cultivation (see Lakela & Craighead; Morton, 1976; Poppleton et a/.; Small; Ward: Wunderlin; and acknowledgments), but neither appears to have become a component of the local flora. In Florida Hedychium coronarium, of the tribe Hedychieae, is more widespread outside of cultivation than the species of A/pinia. Zingiberaceae are characterized vegetatively as mostly terrestrial, rhizoma- tous, sometimes large herbs with aromatic oils in specialized cells (Zingiber- oideae) and with ligulate leaf sheaths. Diversity in overall form disguises a basic floral construction that helps to unify the family: the tubular calyx tends to split along one edge; the corolla and androecium form a tube, from which arise three petals, often two petallike to toothlike (or inconspicuous or absent) lateral staminodes, one fertile stamen bearing a complete anther, and a usually large labellum formed by the fusion of at least two staminodes. The thin style characteristically lies in a groove in the filament and passes between the locules of the anther. The arillate seeds contain straight embryos. Polyploid and aneuploid changes are notable aspects of the evolutionary history of the Zingiberaceae. In fact, according to Sharma & Bhattacharyya, visible changes in chromosome complements have been sufficiently pervasive for different species typically to have different karyotypes. These authors, who studied both wild and cultivated Zingiberaceae, found that chromosomal het- erogeneity is significant even within individuals. A second noteworthy cyto- 1984] ROGERS, ZINGIBERALES 17 logical aspect of the family is that it should probably be counted along with the Juncaceae, Cyperaceae, and Musaceae(?) as one of the few families con- taining species that have chromosomes with diffuse centromeres (Grant). A taxonomic history is available in Holttum’s (1950) partial revision of the Zingiberaceae; historical notes and a compilation of important recent papers appear in Burtt (1972). In a paper rich in information on older literature, Burtt & Smith (1972a) concentrated on 47 ‘“‘key species” in an attack on the snarled early history of the family. Because so much has been written about the Zin- giberaceae, the present historical discussion is limited to selected recent changes and major problems, the most controversial of which may be the choice of rank for the group often called subfam. Costoideae. Subfamily Costoideae is commonly raised to the rank of family by modern authors, particularly those concerned with systems of classification and inter- relationships of families on a broad scale (Cronquist; Dahlgren; Takhtajan; Thorne: and Tomlinson, 1962, 1969). In contrast, others who have worked intensively on the taxonomy within the groups under consideration have rel- egated Costoideae to the subfamilial rank (Burtt & Smith, 1983, Holttum, 1950: Maas, 1972 and subsequent updates), and Hutchinson gave it only tribal status. That differences separating Costoideae from other Zingiberaceae span a wide spectrum of characters helps to counterbalance the fact that many of the dif- ferences are based on small samplings of species. Costoideae stand apart veg- etatively in having strongly developed main stems, spirally arranged leaves, closed leaf sheaths (also true of some Zingiberoideae—see Spearing) bearing ligules of a characteristic nature (Tran van Nam), and no aromatic oil cells. The flowers have nectaries confined within cavities (vs. rising into the floral tube), may differ in anatomical details (Rao et a/.), and lack apparent lateral staminodes, which is also true of some Alpinieae, and which is possibly due to fusion with the labellum. Embryologically, Costoideae may be distinguish- able by the nature of the nucellus, by having a tetrasporic (vs. monosporic) megagametophyte (Mauritzon) and an expanded cotyledon (vs. being haus- torial—see Weisse, 1932), and by additional characters. Pollen of Costoideae is usually panto- or diaperturate (vs. mostly inaperturate) and has thin to thick exines (cf. Hedvchium, Canna—see Erdtman; Maas, 1972; Punt). The two groups are further separated by differences in anatomy (Tomlinson, 1956, 1962. 1969), flavonoids (Williams & Harborne), and centers of distribution. Most, but not all, chromosome numbers in Costoideae are multiples of nine, but the broadly ranging numbers in Zingiberoideae also include multiples of nine. Raghavan & Venkatasubban’s assertion that chromosomal morphology 1S another distinction was not borne out by Sharma & Bhattacharyya. Beyond the evident need for confirmatory surveys, reluctance of some au- thorities to elevate Costoideae to the rank of family may rest partly upon the emphasis given technical and vegetative characters versus that given the fun- damentally “‘zingiberaceous” flower structure that links the two groups. Flower structure is traditionally a weighty character for delineating families of Zingi- erales. Schumann (1904 and earlier) broke subfam. Zingiberoideae into three tribes 18 JOURNAL OF THE ARNOLD ARBORETUM VOL. 63 differing primarily in the number of locules in the ovary, the nature of the placenta, and the form of the lateral staminodes. Holttum (1950) transferred Zingiber from Schumann’s Zingibereae (and consequently changed the name of this tribe to Alpinieae) to the Hedychieae, which should be renamed Zin- gibereae if this transfer is accepted. (Cytological support given by Mahanty for the transfer was dismissed as unsound by Burtt & Olatunji; better corroboration is provided by the plane of distichy of the leaves.) In 1972 Burtt & Olatunji placed Zingiber in a tribe of its own, thereby increasing the total of tribes to four: Alpinieae Meisner, Zingibereae (beware of disparate applications of this name), Hedychieae Duchartre, and Globbeae Meisner.° Dissatished with tribal delimitations, Burtt (1972; see also Burtt & Smith, 1972b; Smith, 1980a) dis- cussed limitations on the taxonomic value of placentation and the nature of the lateral staminodes and agreed with Weisse (1933) that, although continued surveys are needed, the transverse plane of distichy on the aerial shoot in relation to the rhizome may distinguish Alpinieae from Hedychieae, in which the plane is parallel (as in Zingibereae sensu Burtt & Olatunji and those Glob- beae examined by Weisse). Zingiberopsis Hickey, a genus of three species of fossil leaves dating back to the Late Cretaceous and known from Alberta, North Dakota, Wyoming, and Colorado, was placed in the Zingiberaceae by Hickey & Peterson. Widespread in Eurasian sediments of Eocene to Pliocene age, zingiberaceous trilocular fruits and arillate seeds of Spirematospermum Wetzleri (Heer) Chandler resemble those of the modern Asian Cenolophon oxymitrum (K. Schum.) Holttum (Friedrich & Koch, 1970, 1972 Chemical information on Zingiberaceae is scanty in relation to the size of the family. The summary that follows was condensed chiefly from surveys in Gibbs, Williams & Harborne, and Hegnauer. The aromatic oils of Zingiber- oideae are rich in monoterpenoids (among them linalool, camphene, pinene. sabinene, borneol, camphor, cineole, and many others—the last three, accord- ing to Hegnauer, often the chief constituents of the oil). Sesquiterpenoids (in- cluding bisabolene, curcumenes, humulene, caryophyllene, zingiberene, tur- merone, atlantones, zerumbone) are common and (hydrocarbons among them) sometimes predominate in the oil. Likewise widespread among aromatic plants, phenylpropane compounds are represented in zingiberaceous oils, sometimes as major components, by cinnamic acid and at least one derivative. Flavonoids are abundant, and Zingiberaceae (a small sample of Zingiberoideae) “form a chemically well defined group in which a variety of common (quercetin and kaempferol) and more unusual (myricetin, isorhamnetin and syringetin) fla- vonols occur in glycosidic combination with glucuronic acid. rhamnose or glucose” (Williams & Harborne, p. 226). Mostly because of their aromatic oils, derivatives of Zingiberaceae have been used since ancient times as spices and condiments, in perfumes, and medici- nally. Among the many products obtained from the family are ginger (prin- — ‘Tribal names used here differ from those in Burtt (1972) in the spelling of Alpinieae and in the changed authorship of Hedychieae to reflect its valid publication by Duchartre (Dict. Hist. Nat. 13: 849.) 1984] ROGERS, ZINGIBERALES eed cipally rhizomes of Zingiber officinale Roscoe); turmeric (rhizomes of Curcuma longa L. (C. domestica Valeton), an ingredient of curry powders and the source of a yellow dye used like the more expensive saffron); melegueta pepper (seeds of Aframomum Melegueta (Roscoe) K. Schum. ); cardamom (capsules and seeds of Elettaria cardamomum (L.) Maton and substitutes from other species); and galangal (A/pinia officinarum, A. Galanga (L.) Willd.). An “arrowroot” starch is extracted from rhizomes of Curcuma angustifolia Roxb. and other species. Some of the many genera grown as ornamentals under warm conditions are Alpinia, Hedychium, Kaempferia L., Nicolaia Horan. (Phaeomeria Lindley ex K. Schum.), and Roscoea James Sm. Species from a number of genera have been used as sources of fibers or pulp. (Among several works dealing with economic aspects of Zingiberaceae are Burtt, 1977a, 1977b, 1980; Ilyas; Mor- ton, 1981; Perry; Purseglove; Rosengarten; and Schumann.) REFERENCES: Under ordinal references, see CostTeRus (1916b); CRoNQuist (1981); Czasaz DAGHLIAN: DAHLGREN; Davis; ERDTMAN; Gibps; HEGNAUER; HoRANINOW;, HUMPHREY; HutTCHINSON:; LAKELA & CRAIGHEAD; MAHANTY; MAURITZON; MORTON; OLATUNJI, PER- RY: POppLETON et al., PoTZTAL: PURSEGLOVE:; A. S. RAO & VERMA; V. S. Rao, V.S. Rao & Gupte: V. S. Rao, KARNIK & Gupte; V. S. RAo & PAI, RICKETT: Rosco OE; SATO; SCHACHNER: SHARMA: SMALL. TAKHTAJAN: THORNE; TOMLINSON; VENKATASUBBAN; WARD: WILLIAMS & HARBORNE; and WUNDERLIN. Burtt, B. L. General introduction to papers on Zingiberaceae. Notes Bot. Gard. Edin- burgh 31: 155-165. 1972. pee list of subfamilies and tribes with corrected authorship, citations, and synony . Curcuma Zedoaria. Gard. Bull eee 30: 59-62. J unnumbered pl. 1977a. [C. Zedoaria (Christm.) Roscoe.] _ The nomenclature of turmeric and other Ceylon Zingiberaceae. Notes Bot. Gard. Edinburgh 35: 209-215. 1 Cardamoms and other Zingiberaceae in Hortus Malabaricus. Pp. 139-148 in K. S. MANILAL, ed., Botany and history of Hortus Malabaricus. Rotterdam. 1980. O. A. OLATUNIL. re limits of the tribe Zingibereae. Notes Bot. Gard. Edin- burgh 31: 167-169. 1972. &R SmitH. Key species in the taxonomic history of Zingiberaceae. Notes Bot. Gard, Edinburgh 31: 177-227. 1972a. [Hedychium coronarium, 197, 200: see later papers of he + for updated nomenclature for some spec , ntative keys to the subfamilies, tribes and ee of Zingiberaceae. Ibid. 171-176. ‘1972, ngiberaceac. Pp. 488-532 in M. D. DassANAYAKE & F.R, FOsSBERG, eds., A revised eee to the flora of Ceylon. Vol. 4. New Delhi. 1983 CHAKRAVoRTI, A. K. Multiplication of chromosome numbers in relation to speciation in Zingiberaceae. Sci. Cult. 14: 137-140. 1948.* . Cytogenetical studies in Zingiberaceae. Proc. Indian Sci. Congr. 39(3): 30, 31. 1952. [This and the preceding reference a interest in connection with diffuse cen- tromeres in Zingiberaceae; also see GRAN FriepRicH, W. L., & B. E. Kocnu. Ca of fruits and seeds of fossil Spiremato- spermum (Zingiberaceae) with those of living Cenolophon. Medd. Dansk Geol. For. -195. pl. 1. 1970. [Fruits and seeds of Miocene age from Denmark; also see DAGHLIAN (ordinal references).] & Der Arillus der tertidren Zingiberacee Spirematospermum Wetzleri. (English abstract.) Lethaia 5: 47-59. 1972. [The affinity suggested in the preceding 20 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 paper buttressed by SEM observations on arils; ca. 100 seeds and some fruits from 6 European localities. ] Furterer, W. Beitrage zur Anatomie und Entwicklungsgeschichte der ere Bot. Centralbl. 68: 241-248, 273-279, 346-356, 393-400, 417-431. 10, 35-46. 1897. [Species of Hedychium, Brachychilum, Kaempferia, poate Cur- cuma, Costus, Alpinia, Zingiber, and Globba. GRANT, V. Plant speciation. ed. 2. x11 + 563 pp. New York. 1981. [Zingiberaceae, 370; evidently feels that confirmatory work is needed to establish occurrence of diffuse centromeres in Zingiberaceae: see also CHAKRAVORTI. GROOTIEN, C.J. & F. BouMAN. Development of the ovule and seed in Costus cuspidatus N. E. Br.) Maas (Zingiberaceae), with special reference to the formation of the operculum. Bot. Jour. Linn. Soc. London 83: 27-40. 1981. [Includes table comparing families of Zingiberales for 3 characters of seeds.] Hate, N. Architecture du rhizome chez quelques Zingibéracées d’ Afrique et d’Océanie. Adansonia, II. 19: 127-144. 1979. [Seven genera.] Hickey, L. J... & R.K. Peterson. Zingiberopsis, a fossil genus of the ginger family from Late Cretaceous to early Eocene sediments of western interior North America. pes Jour. Bot. 56: 1136-1152. 1978. [Cf DAGHLIAN (ordinal references). ] Hoitrum, R. E. The Zingiberaceae of the ue Peninsula. Gard. Bull. Singapore 13: “250. figs. 1-33 on 33 unnumbered pages. 1950. [Includes details of floral biology, morphology, taxonomic history, Soeacre and revised subfamilial and tribal arrangements. A commentary on comparative morphology in Zingiberaceae. Ibid. 27: 155- 166. 1974. {Includes illustrated discussion of the architecture of the inflorescence and advocates more taxonomic use of inflorescence. } Inyvas, M. The spices of India—II. Econ. Bot. 32: 238-263. 1978 [1979]. [Diverse ei on a number of zingiberaceous spices, including lists of compounds as ted in their oils and a bibliography with references not included in the present ne Kumar AR, V. Course . vascular differentiation in the axis of certain Zingiberaceae. Flora 162: 420-425. 1973a. [Two species of € urcuma.| : Raat rea studies in the roots of Zingiberaceae. [hid. 467-478. 1973b. Lorsenrr, T. Zingiberaceae. Nat. Pllanzenfam. ae 2. 15a: 541-640. 1930. Maas, P. J. M. Costoideae erg : ne otrop. Monogr. 8: 1-140. 1972. [Updated in Acta Bot. Neerl. ae 975: Fl. Neo oe ee 18: 162- 209, 1977; and in the ae oo : Costus, x = 9, 2n = 28 reported in Dimerocostus and rea ] Zingiberaceae. /n. utter ING & B. SPARRE, , Fl. Ecuador 6: 1-50. 1976. MenHra, PN. & S. K. ee HDEVA, tological eran on some W. Himalayan monocots. IV. Several families. : re 41: 31-53. 1976. [Zingiberaceae, 34. 47- 513) PANCHAKSHARAPPA, M. G. Taxonomic evaluation of Zingiberaceae. Bull. Bot. Surv. India 4: 129-135. 1962 [1963 Punt, W. Pollen duet! of the American vibee of the subfamily Costoideae (Zingiberaceae). Rev. Pa Palynol. 7: 31-43. 1968. RAGHAVAN, T. S., & K. R. NKATASUBBAN, ee logical studies in the family Zingi- beraceae with special ae to chromosome number and cyto-taxonomy. Proc. Indian Acad. Sei. 17B: 118-132. 1943. [Chromosome counts for 24 species {ro enera, chromosome numbers from other literature. and details of cnromasoinel morphology. | RAMACHANDRAN, K, Chromosome numbers in Zingiberaceae. Cytologia 34: 213-2 . (Ca. 26 spp. from 11 genera; includes chromosome numbers from othe oan and photographs of chromosomes of MHedyvchium coronarium. ROSENGARTEN, F. The book of spices. x11 + 489 pp. Wynnewood. Pennsylvania. 1969. (Cardamom, 160-173: ginger, 254-265: turmeric, 444-452. ] 1984] ROGERS, ZINGIBERALES ra\ Royen, P. vAN. The alpine flora of New Guinea. Vol. 2: taxonomic part, Cupressaceae to Poaceae. Ixvili + 1232 pp. + addendum. a Liechtenstein. 1979. [Zingiber- SAC HA ARS R. Oe ial U. ARORA. Some a ous aspects of Amomum dealbatum and natum. Bot. Gaz. 124: 353-360. 1963. [Includes gener See n fa mily prone by brief review of literature, photo of sectioned seed of Hedychium, illustrated embryogeny and development of megagametophyte, and description of polyembryony in Hedychium.] SCHUMANN, K. Monographie der eee von Malaisien und Papuasien. Bot. Jahrb. 27: 259-350. pls. 2-6. [1899] | Zingiberaceae. Pflanzenr. IV. pene 20): 1-458. 1904. [See criticism of this monograph in Hotrrum (1950), pp. 16, 17.] SHarMa. A. K.. & N. K. BHATTACHARYYA. Cytology of several members of Zingiberaceae and a study of the inconstancy of their chromosome complements. Cellule 59: 297- 346. pls. 1-3. 1959. [Includes table of chromosome counts for the entire family from other literature plus original observations; another table shows chromosomal mor- phologies of species eee: several genera, among them 8 species of He- dychium, in the family 2” = 18-62 SHEEHAN, T. J. Zi basen for Florida. Proc. Florida State Hort. Soc. 71: 382-388. 1958.* > = Smitu, R. M. Anew genus of Zingiberaceae from N Burma. Notes Bot. Gard. Edinburgh 38: 13-17. 1980a. [Stadiochilus, comments on tribal arrangement in Zingibera ceae. | _ Notes on Zingiberaceae. Ibid. 19-21. 1980b. [Includes Hedychium tenellum (K. Schum.) R. M. Sm., comb. nov.] SPEARING, J. K. A note on closed leaf-sheaths in Zingiberaceae-Zingiberoideae. [bid. §: 217-220. pl. 2. 1977. [Closed sheaths in ules and Cautleva.] Steiner, M. L. A new and illustrated flora of Manila, I. Zingiberaceae. Philip. Jour. Sci. 88: 1-40. pis. "3: 1959 [1960]. [Eleven genera, including Hedychium (2 intro- duced species); mentions an additional species of Hedychium indigenous to the Philippines. ] TOML hee P. B. Studies in the systematic anatomy of the Zingiberaceae. Jour. Linn. Soc. Bot. 55: 547-592. 1956. [Despite importance in distinguishing subfamilies and ea at the generic level, anatomical data of limited value at the tribal level.] Tran van Nam. Costaceae et Zingiberaceae: leurs appareils ligulaires. Adansonia, II. 14: 561-570. 1975. [Believes that ontogenesis and tracheogenesis of the ligules confirm that these groups are distinct families; ligule of Zingiberoideae (-aceae) appears to differ from that of Costoideae (-aceae) in the pattern of venation and in being of epidermal (vs. subepidermal) origin. ] Weser, A. Die Homologie des Perigons der Zingiberaceen. Ein Beitrag zur Morphologie und Phylogenie des Monokotylen- aacoe (English abstract.) Pl. Syst. Evol. 133: 149-179. 1980. [Studies on a species of Nicolaia help to demonstrate that tepals of Zingiberaceae correspond to leaf ion + “Oberblatt Weisse, A. Zur Kenntnis der Blattstellungsverhaltnisse bei den Zingiberaceen. ae Deutsch. Bot. Ges. 50a: 327-366. pls. 8, 9. 1932. [Foliar arrangements in Costoidea and Zingiberoideae succinctly compared, 361-363, seedling of Hedychium coro- narium described. Die Art der Distichie an den Achselsprossen von Zingiber. Ibid. 51: 13-20. 1933, 1. Hedychium J. G. Koenig in A. J. Retzius, Obs. Bot. 3: 61 (°73”). Lees. Upright, often large (17. coronarium to 3 m tall), [frequently epiphytic] herbs, the unbranched, leafy aerial stems borne on horizontal rhizomes and dying after flowering. Leaf blade inserted immediately above the sheathing base [or 22 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 on a short petiole], usually attenuate at both ends, sometimes bearing soft. pale, matted hairs abaxially. Inflorescences usually spicate [rarely branched]. with large, crowded, spirally arranged [or reportedly sometimes whorled], im- bricate bracts covering the axes, each bract usually subtending multiple flowers [or the bracts divergent from the then exposed axis. enfolding | or more flowers]. Flower bracteolate, fragrant, white and often with yellow toward the base of the labellum and the bases of the lateral staminodes [or colored differently, the colors then usually yellows to reds or sometimes purple]. Calyx tube short, split along | side [or entire], 3-lobed to apparently |-lobed, usually glabrous with indument concentrated at the apex (//. coronarium). Corolla tube long and slender, projecting beyond [or shorter than] the subtending bract, much longer than [or infrequently about as long as] the calyx. Lateral staminodes 2 broad, petaloid and showy; labellum, the broadest and showiest component of the flower, abruptly [or gradually] narrowed at base, usually apically cleft, the depth of apical notch, width, and overall shape highly variable, more or less obcordate in H. coronarium [to Y-shaped, wedge-shaped, or elliptic]; filament about as long as the labellum, sometimes Slightly shorter [to much longer]: pollen grains spherical, inaperturate or uniaperturate (in //. coronarium the aperture, when present, ulceroid a not clear in surface view, fide Saa Ibrahim). Ovary 3-locular, with many ovules per locule. glabrous to sericeous, the style filiform, lying in a groove in the corolla tube and filament, and pro- truding apically beyond the anther. Capsule loculicidal, 3-locular (fruiting spec- imens not seen from the United States). Seeds variably shaped, with initially red (becoming yellowish) arils, these lacerate on mature seeds. Base chromo- some number mostly 17. Type species: //. coronarium J. G. Koenig.° (Name from Greek, hedvs, sweet, and chion, snow.) _~ A genus of approximately 50 to 60 species, about a third of them described since Schumann’s (1904) revision and many unclearly delimited, in two sub- genera, HEDYCHIUM and EvosMIANTHUS K. Schum.’ The largely Himalayan center of distribution encompasses northeastern India, Bangladesh, the upper Ganges River, and Nepal. Continuing to the south and east with a diminished number of species, the probable ae range of Hedychium, outlined by southern China, Vietnam, the Philippines, Java, and Sulawesi, crosses Wal- lace’s line but does not reach New cae or Australia. At the western extreme, ‘Burtt & Smith (1972a, p. 190) have argued against rejection of this and others of Koenig’s specific names due to publication in a work in which Linnaean binomials were not consistently used. 1853) and Horaninow (1862) each recogniz zed four pine of unspecified rank. two of which remain of interest: “Gandasulium Horan.” and *Macrostemium Horan..” adopted by Baker (1892) at the rank of section and distinguished by the rélalive lengths of the stamen and labellum, Later, Schumann (1904), employing characters of the inflorescence and its bracts, divided the genus Into “subgen. oo Horan.” and subg. Euosmianthus K. Schum. Lourteig (1972) changed the name of the former to subg. //edychium, since it includes the type species of the genus, and adopted for the i eae. Macrostemuun Horan. (sensu Euosmianthus Schum.).” with 7 ~ 2 > Gardnerianum Ww ae ex Roscoe as SUSELOLY De: cis Caer than Luosmianthus, Macrostemiun vas published ICBN, 1978, lacks nomenclatural priority at hie rank of Suse It seems reasonable. neverihels ss. to maintain ff. Gardnerianuim as lectotype for subg. Euosmiuanthus K. Schu 1984] ROGERS, ZINGIBERALES 23 Hedychium peregrinum N. E. Br. is endemic to Madagascar. The natural dis- tribution of H. coccineum Buch.-Ham. ex James Sm. doubtfully includes Sri Lanka (Burtt & Smith, 1983). Species of this widely cultivated genus are scat- tered in other regions largely, if not completely, as a result of human activity. Hedychium coronarium grows extensively outside of cultivation in warm re- gions worldwide. Habitats of species of Hedychium range from tropical to temperate, at altitudes approaching 3000 m, in usually (but not always) open, typically wet places: often shores, moist slopes, and edges of forests. Several species are epiphytic or facultatively so. Subgenus HepycHiuM (subg. Gandasulium (Horan.) K. Schum.) (inflores- cence compact, usually broadly ellipsoid or ovate in outline, with wide, im- bricate bracts covering the rachis) is represented in the southeastern United States by H. coronarium, ginger-lily, butterfly-lily, garland flower, which has escaped cultivation into various wet habitats in Florida (and according to Small, but not substantiated in recent checklists, in eastern Georgia and southeastern Louisiana). Although obscured by cultivation, the natural range of H. coro- narium probably lies within the generic center of distribution described above. In that region H. coronarium is fertile (see, for example, Roxburgh, Mukherjee), although in some or all of the regions into which it has been introduced, production of fruit appears to be rare. Harling thought deficiency of pollinators to be a more likely explanation for low fertility of this species in Ecuador than incompatibility between members of the probably highly clonal population there, Self-incompatibility, however, has been reported in Hedychium (Holt- tum, 1950), and the plant illustrated in Ficure | failed to set fruit with its own pollen. These reproductive limitations notwithstanding, H. coronarium can be an aggressive invader aided by strong rhizomatous growth and possibly dis- tributed by fragmentation and flotation of rhizomes. It forms dense clumps or extensive colonies, in tropical America covering large areas and reportedly sometimes having a deceptively indigenous appearance (Beadle, Standley & Steyermark). The name “Hedychium coronarium’ is applied to various portions of a problematic complex that also involves the names H. maximum Roscoe, [7 flavum Roxb. (non Roscoe), H. flavescens, H. chrysoleucum Hooker, H. uro- phyllum Lodd., H. Elwesii Baker, and 77. subditum Turrill (77. flavum Roscoe). Baker treated most of these as varieties of H. coronarium separated from each other primarily by the coloration of the flowers. Emphasizing this character and the shape of the labellum, Schumann (1904) elevated most to specific rank, as did Turrill in a thorough, illustrated discussion of the matter. In Turrill’s paper an expanded set of vegetative and reproductive characters augments the persistently important floral coloration. Studying living plants, Naik & Pani- grahi uncovered intergradation in the colors of flowers among the species; they preferred using the structure of the inflorescence and the form of the labellum for taxonomic purposes and reassembled H. coronarium in its broader sense. Orchard, Lourteig, and Burtt & Smith (1983) have separated H. flavescens from H. coronarium at the specific level. If judged from the works cited above, je8 coronarium sensu stricto is characterized by having the flower purely white or tinged with yellow or green, the calyx glabrous or nearly so and proportionately 24 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Ficure |. Hedychium. a-n, //. coronarium: a, unbranched stem rising from hori- zontal rhizome, = ',: b, portion of stem showing ligulate leaf bases, * 2: c, cross section of stem, x 6; d, detail of leaf showing venation, x . e, cross section of leaf sheath, x 6; f, inflorescence, x 12; g, flower, showing split, -lobed calyx, narrow petals with incurled margins, 2 broad lateral staminodes, label fee with bifid apex, and single sta- 1984] ROGERS, ZINGIBERALES Z short, the labellum about as wide as long, and the white staminal filament either about as long as or shorter than the labellum. The complex as a whole is recognized by having each bract with multiple mostly white to yellow flowers, each with a large labellum and usually broad lateral staminodes. Asa genus, hedychiums are robust rhizomatous herbs with unbranched aerial stems bearing terminal, usually spicate, many-flowered inflorescences. The fragrant flowers, usually a few per bract, tend to have lopsided calyces usually split along one side; each flower has a longish, narrowly cylindrical corolla tube with none (vs. one in Odontychium K. Schum.) of the lobes appendiculate; two strongly developed (vs. inconspicuous in 4/pinia), petallike lateral staminodes; a protruding (vs. very short in Brac hychilum (R. Br. ex Wal 1.) Petersen), usually bi leaf flavonoid profiles,” Williams & Harborne (p. 224) pointed out that my- ricetin glycosides are the predominant flavonoids in Hedychium (9 species studied). Smith (1980a) found the new genus Stadiochilus R. M. Sm. to resemble Hedychium in having the corolla tube grooved to accommodate the style (known also in Brachychilum and Zingiber), in the form of the corolla lobes, and in having a long filament. Smith (p. 14) stated that “if a pair of petaloid lateral staminodes were added to Stadiochilus we should have a perfectly good He- dychium rendered slightly anomalous by the upright, rather pendulous label- lum.” Most chromosome counts in Hedychium have yielded the somatic number 34, although the genus appears to be prone to polyploidy and aneuploidy, with numbers varying even within species (see especially Mukherjee). The following somatic numbers have been reported in species of Hedychium: 18 (H. coro- narium, see Hsu), 24 (reported once in H. thyrsiforme Ham., but contradicted by further study—see Mahanty), 26, 36, 50, 51, 52, 54, 66, and 68. Most of these numbers up to 66 have been reported in H. coronarium sensu lato, as well as in other species. Raghavan & Venkatasubban observed general simi- larity in the size and morphology of chromosomes among the six(?) species of Hedychium that they studied. Similarly, although the chromosomal morphol- ogy was not identical in any two taxa, Sharma & Bhattacharyya noted ‘‘a gross resemblance’”’ among 10 species and varieties of Hedychium. Those two au- thors, in good agreement with Mukherjee, found the chromosomes of He- men, < '2; h, adaxial side of anther (note thin style passing between the 2 locules, the se protruding apically), x 3;1, abaxial view of anther, x 3; J, lateral view of stigma, 5: k, adaxial view of stigma, x 15; 1, diagrammatic cross section of corolla mie nie! calyx, the lumen in black, the style in groove above lumen unshaded, x diagrammatic cross section of flower slightly above ovary, showing calyx (outer unshaded layer not adnate to corolla tube), corolla tube (unshaded), 2 epigynous nectar glands, and style (in groove in corolla tube), x 6; n, diagrammatic cross section of ovary, showing locules and axile placentae, x 6. 26 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 dychium generally to be short, varying graduall y in length within each com- plement. The different potential species of Hedychium studied by Sharma & Bhattacharyya had chromosomes with the primary constrictions nearly sub- median to median and varied in the numbers of pairs bearing secondary con- strictions. Mukherjee and Sharma & Bhattacharyya reported meiosis in H. coronarium to be mostly normal or nearly so; Mukherjee recorded 96 percent pollen fertility in material from northeastern India. According to Holttum (1950), hybrids are easily produced in /Hedychium. The three main layers of the walls of pollen grains in //edvchium resemble those of the more thoroughly studied Canna and certain other genera. In cea al a eta ea thin, acetolysis-resistant, presumably exinous layer overlies a thick, acetolysis-degradable, almost certainly intinous layer with a adie: patterned infrastructure. The third, innermost layer is hygro- scopic and lacks a radial pattern (Erdtman & Praglowski). Although generaliza- tions would be premature and substantial variations have turned up, there are indications that pollen roughly as described above may be widespread among Zingiberales (see Saad & Ibrahim; Erdtman, 1963; Hesse & Waha, under Can- na: Kress et al.. Kress & Stone, under Canna). Meager data point to sphingids as the predominant pollinators of Hedychium coronarium and probably most other species of the genus. Pollen may some- times be transferred on the wings of butterflies (Miller). Accessory embryos were observed to arise from synergids but not to develop beyond early stages in H. acuminatum Roscoe (Sachar & Arora). Hedychium coronarium, H. coccineum, H. Gardnerianum Roscoe, H. spt- catum Buch.-Ham. ex James Sm. (see Winters & Corbett) and several other species are cultivated, mostly under warm conditions. The fragrant, showy flowers of H. coronarium are valued for making Hawaian leis (see Neal). Vol- atile oils from flowers and rhizomes of this and possibly other species are useful in perfumery. Rhizomes of //. spicatuim, sometimes sold as the drug “kapur(a) kachari’ and sometimes in a fragrant powder, abir, lend their scent to tobacco, incense, soaps, face powders, hair oils, and fabrics, and they are thought to repel insects. Medicinal uses of derivatives of H. spicatum and other he- dychiums are available in Chaturvedi & Sharma, Cooke, Dixit & Varma, Perry, Pineda-Ocampo et al., Sastri, B. D. Sharma (1974, 1975), and the papers by S. C. Sharma et al. Hedychium coronarium has attracted attention as a rapidly growing source of pulp for paper. This rhizomatous species sometimes becomes a weedy pest in places where sugar cane Is grown (Beadle REFERENCES: Under ordinal references, see DUNCAN & Karresz, HORANINOW, MAHANTY, MULLER, Near, Perry, Rickett, SAAD & IBRAHIM, SMALL, SrANDLEY & STEYERMARK, THIERET. and WitiamMs & Harborne; also see ordinal references listed at beginning of family references; under family references, see BURTT & SMITH (1972a, 1983), Hotrrum (1950), RAGHAVAN & VENKATASUBBAN, SACHAR & ARORA, SCHUMANN, SHARMA & BHATTACHARYYA, SMitH, and Weisse (1932). ANONYMOUS. New sources of paper aera coronarium, Koen., and allies). Bull. Misc. Inf. Kew 1912: 373-378. pls. 1, 912 1984] ROGERS, ZINGIBERALES pe —. Hedychium coronarium from Calcutta. /bid. 1914: 165-167. 1914a. [See also p. 175.] —. Hedychium flavescens. Ibid. 1914: 193, 194. 1914b. [Compared with H. coro- narium as a source of paper. BAKER, J. G. Hedychium. In: J. D. Hooker, FI. ne India 6: 225-233. 1892. BAtuie, H. P. peLA. Zingibéracées, Fam. no. 47. Jn: H. Humbert, Fl. Madagascar et des Comores. 34 pp. 1946. [Hedychium, 4- en H. coronarium var. flavescens (es- caped); H. peregrinum (endemic); see also LouRTEIG.] BeaAp_e, C. Hedychium coronarium in Brazil. Bull. Misc. Inf. Kew 1917: 104, 105. 1917. [Description of stands and comments on distribution CuHaturvept, G. N., & B. D. SHARMA, Clinical studies on Hedychium ee ail an anti-asthmatic drug. Jour. Res. Indian Med. 10: 6-10. 1975.* [Abstract in Ex- cerpta Bot. 30: 124. 1977; drug decreased respiratory rate and absolute sere count and increased the vital capacity in patients with bronchial asthma: see also B. D. SHARMA (1975).] Cooke, M.C. Kafur kachri (Hedychium spicatum). Pharm. Jour. II. 1: 603, 604. 1871. [Sold in India as a drug, considered to be a tonic and stimulant, used to scent tobacco; includes illustration of starch eee : Dixit, V.K.,& K.C. Varma. Anti-helminthic es of essential oils from rhizomes of Hedy en coronarium and ene cena Indian Jour. Pharm. 37: 143, 144. 1975 _N. VaAsHISHT. Studies on essential oils of rhizomes of Hedychium spicatum Koenig and Hedychium coronarium Koenig. Indian Jour. Pharm. 39: 58- 0. 1977. [Isolated from rhizomes: 6-caryophyllene, 8-caryophyllene oxide, linalool, elemol, a-pinene, 8-pinene, limonene, A?-carene, 6-phellandrene, p-cymene, and cineole (this most abundant). ] ErptMaAn, G. Sporomorphology and phytomorphology. Jour. Indian Bot. Soc. 42A: 35- 38. pls. 1, 2. 1963. [Pl 1, fig. 4, section through part of a pollen grain from H. coronarium. | & J. R. PRAGLOwskI. Six notes on pollen morphology and pollen-morphological techniques. Bot. Not. 112: 175-184. pls. 1-6. 1959. [H. coronarium, 178, 179, pl. 6 (sectioned pollen grain.] Fano, D. Some new taxa of Zingiberaceae from Kwangsi. Acta Phytotax. Sinica 16(3): 47-53. 1978. [H. tienlinense D. Fang, sp. nov., 52, 53. Hara, H. The flora of Eastern Himalaya. xi + 744 pp. + 40 pls. + map. Tokyo. 1966. [Hedychium, 422.] HarinGc, G. Zur Embryologie der Gattung Hedychium Koenig. Sv. Bot. Tidskr. 43: 357-364. 1949. [If deficient fruit production is due to failure of pollination, endo- sperm must develop without pollination; endosperm helobial; embryo sac of the Polygonum type. Hsu, C. C. Preliminary chromosome studies on the vascular plants of Taiwan (1). Taiwania 13: 117-129. 1967. [#f. coronarium, 128. Kress, W. J., D. E. Stone, & S.C. SELLERS. oe of exine-less pollen: Heliconia (Heliconaceae, Am. Jour. Bot. 65: 1064— ae 78. KUNCKEL D’HercuLAaIs, J. Rapport des insectes L ke avec les fleurs des Zin- gibéracées et en particulier avec celles des fea. Compt. Rend. Acad. Sci. Paris 151: 1153-1155. 1910. [Describes cases in which certain Sphingidae unable to disengage themselves from floral tube die attached to flower.] Larsen, K. Studies in Zingiberaceae V. Hedychiuim—some new Indo-chinese taxa. Bot Tidsskr. 61: 71-79. 1965. [H. boloveniorum, H. Poilanit, H. Forrestii var. latebrac- teatum, all new.] Lourteic, A. Le genre Hedychium a Madagascar (Zingibéracées). Adansonia, I. 12: 21-127. 1972. Mapce, M. Nuclear migrations in Hedychium. Proc. Linn. Soc. London 146: 108, 109. 28 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 1934. [In H. Gardnerianum nucellar nuclei reported to migrate from cell to cell; see comments in HARLING.] Division of the generative cell in Hedychium eae Cellule 45: 169- 176. 1 unnumbered pl. 1936. [Describes growth of polle Mipatey, J. B. Hedychiums for glasshouse culture. Gard. oe Ill. 170: 36. 1971. MuknHerJeEE, I. Chromosome studies of some species of Hedychium. Bot. Mag. Tokyo 83: 237-241. 1970. [x = 17; polyploidy and aneuploidy common within and between species; 7 species, including H. coronarium. mate oe list of chromosome numbers from other literature and drawings of chromosomes. ] Naik, V. N., & G. PANIGRAHI. Genus Hedychium i in eastern India. Bull. Bot. Surv. India 3: 67-73. 1961. [Generic description, key to species, taxonomic history of /7/. coronarium complex, discussion of problems connected with other species, enu- meration of 17 species with distributions and short descriptions. ] OrcHarb, A. E. Hedychium in New Zealand—a further name change. New Zealand Jour. Bot. 16: 287-289. 1978. [Species called H. flavum Roscoe should be called /1. flavescens Carey ex Roscoe; includes taxonomic history of H. coronarium complex.] PINEDA-Ocampo, M., L. OLIVEROS-BELARDO, & E. Fermin-Sitva. A study of the a a of the rhizome of aes coronarium Koenig. Proc. 8th Pacific Sci. Con : 145-160. 1954. [Includes photo of portions of rhizomes, review of medicinal uses, and description Ro properties of oil; cineole and (probably) camphene detected: oil has depressant effect on cat intestine and carminative effect on humans. } Rao, A.S.,& P. K. Hasna. Hedychium radiatum & H. robustum (Zingiberaceae)— two new species from oe District, Arunachal Pradesh. Bull. Bot. Surv. India 16: 163- ie 1974 9 77). . RMA. Notes on Soa Koenig, including four new species from mere Hills, Assam. Bull. Bot. Surv. India 11: 120-128. 1969 [1971]. [New species: H. calcaratum, H. co ees H. deg ies H. rubrum.| RoxBuURGH, W. Flora Indica. Vol. 1. 11 + pp. Serampore, India, & London. 1832. [Hedychium, 10-14; aah descriptions of seeds and ee of rH. coronarium.| Sastri, B. N., chief ed. The wealth of India. Raw materials. Vol. 5. xxv + 332 + xu pp. New Delhi. 1959, [Hedvehium, 11-14: discussion of H. eee as a source of pulp for paper; many other uses for this and other species; principal component of oil from rhizome of H. spicatum is ethyl ester of p-methoxy cinnamic acid; several other compounds isolated from this species and from //. flavuim.] = B. D. Studies on “kapura kachari” (Hedychium spicatum). Jour. Res. Indian Med. 9: 69, 70. 1974.* [Abstract in Excerpta Bot. 27: 445. 1976; drug vasodilatory on coronary vessels, mildly hypotensive, antispasmodic on smoot musc ou macological evaluation of thet S of Hedye ie spicatum (Banhalthi). /bid. 10: 17-22. 1975.* ee in oe Bot. 30: 164. 1978: anti-asthmatic effect due to anti-histaminic property.] SHARMA, M. Pl morphology of Indian monocotyledons. Jour. Palynol. Special Vol- ume. ii + 95 pp. + 3 charts. 1968. [Hedychium, 10; 6 species ate includes brief generic pls nological characterization. SHARMA, 8. C., Y. N. SHUKLA, & J. S. TANDON. Alkaloids and terpenoids of Ancistro- cladus hey neanus, Sagittaria sagittifolia, Lyonia formosa and Hedychium spicatum. Phytochemistry 14: 578, 579. 1975. [Extract from rhizome of HH. spicatum ee flammatory; contains sitosterol, sitosterol-8-D-glucoside, and cryptomeridio J.S. TANDoN, & M. M. Duar. 7-hydroxyhedychenone, a fura ee from ee spicatum. Phytochemistry 15: 827, 828. 1976. [Determination of struc- e.] . Uprety, Y. N. SH A, & M . Duar. Hedychenone: a furanoid drempent fom Hedychium pees Serves 14: 1059-1061. 1975. [Rhi- zomes used as insect repellent, as perfume for tobacco, and in treatment of stomach ailments and inflammation. ] 1984] ROGERS, ZINGIBERALES ay Sims, J. Hedychium coronarium. Sweet-scented garland-flower. Curtis’s Bot. Mag. 19: pl. 708. 1803. Turritt, W. B. Hedychium coronarium and allied species. Bull. Misc. Inf. Kew 1914: 368-372. 2 unnumbered pls. 1914. [Includes description of H. coronarium, are trations of flowers of this and presumably closely related species, Sea of H. coronarium with other species, instructions for collecting hedychium Watticu, N. Initiatory attempt to define the species of Hedychium, and settle their synonymy. Jour. Bot. Kew Misc. 5: 321-329, 367-377. 1853. [Twenty-three species sorted into Coronariae, Spicatae, Siphonium, and Brachychilum.] Warp, F. K. Hedychiums. Gard. Chron. HI. 120: 18, 19. 1946. [Most commonly cultivated species are H. coronarium, H. coccineum, H. angustifolium, and H. Ga rd- nerianum. Winters, H. F., & E. G. Corpetr. A cold-hardy ginger lly. Am. Hort. Mag. 44: 129- ” 1965. [H. spicatum flourishes in cultivation as far north as Maryland.] CANNACEAE A. L. Jussieu, Gen. Pl. 62. 1789, ““Cannae,”’ nom. cons. (CANNA FAMILY) Large, usually rhizomatous herbs with unbranched aerial stems. Leaves large, with open, eligulate sheaths. Flower asymmetric, with a staminodial lip (la- bellum) and a variable number of additional petallike staminodes [or none], | fertile, partly petaloid stamen, and | more or less flattened, colorful style rising from a 3-locular ovary with septal nectaries. TyPpE GENUS: Canna L. 1. Canna Linnaeus, Sp. Pl. 1: 1. 1753; Gen. Pl. ed. 5. 1. 1754. Plants often | to several meters tall, often glaucous, mostly glabrous (some- times with an ephemeral, woolly indument on leaves and stem [or inflores- cences]). Rhizomes branching sympodially, sometimes thickened, these and a free; stomata predominantly paracytic. Inflorescences terminal on leafy shoots, spicate or branched, each branch with a 2-keeled prophyll. Flowers showy, in homodromous cincinni of 2 flowers (1 flower of the pair typically not de- veloping, rarely with 3 flowers), each flower or pair subtended by a bract and sometimes with scalelike bracteoles, erect [to pendulous], colors mostly yellows and reds. Sepals 3, free, persistent. lanceolate to elliptic. Petals 3, usually unequal, usually basally connate and adnate to the androecium and style to form a tube, upright or reflexed, deciduous. One stamen fertile and partly petaloid, enfolding the style in the bud, with a marginal anther, this sometimes appearing superficially to be 2-locular (but generally interpreted as a |-locular half-anther); the other androecial members sterile, one of these curled [or straight] forming the labellum, the remaining 3 or 2 [or none in subg. Distemon] mostly large, erect, broadened, and petallike (flowers rarely with a fifth sterile staminode); pollen deposited onto the side of the style in the bud, the pollen grains mostly spherical, usually with small spinules, inaperturate (sometimes with | or 2 pores fide Nair, 1962). Ovary covered with conspicuous, sap-bearing protuberances, each of the 3 locules with numerous anatropous ovules on an 30 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 AS oa KS qs ae = og i nae SS eee SAS 4 Figure 2. Canna. a-q, C. flaccida: a, young aerial shoot coming from underground rhizome, x '4; b, blade and portion of petiole of nearly mature leaf, x ‘4: c, detail of upper part of sheathing petiole—ligule not present. x ‘4; d. inflorescence with open, nocturnal flower and flower bud. both in side view—on open flower note bractlike sepals 1984] ROGERS, ZINGIBERALES 31 axile placenta; style opposite the fertile stamen; stigmatic surface terminal and decurrent along an edge of the style. Capsule globose to oblong-ellipsoid, tu- berculate-bristly, loculicidally dehiscent or the seeds released by breakdown of the pericarp. Seed globose or nearly so, dark colored, hard, opening by a slit, with an arillike tangle of hairs arising from the funiculus, these hairs remaining in the fruit; with abundant nutritive tissue (““chalazosperm”’) derived from the chalazal region; endosperm initially nuclear, reduced to a thin layer in the mature seed. Embryo linear, straight, extending into a cavity at the end of the seed. Radicle and plumule developing apart from the remnant of the seed, connected to it by a cotyledonary appendage. Megagametophyte (embryo sac) essentially of the Polygonum type. Base chromosome number 9. LECTOTYPE species: C. indica L.; see N. L. Britton, Fl. Bermuda, 86. 1918. (Name from Greek Kanna, cane.) A distinctive monogeneric family with a problem-free circumscription, in contrast with its internal taxonomic and nomenclatural disarray. The usually quoted estimate of about 50 species in Canna was approximately halved in the revision by Segeren & Maas, and according to Jiménez (citing personal communication), Maas has since compressed his estimate to only seven species. The genus is centered in the American tropics and subtropics, including the West Indies; species occupying scattered warm areas in the Old World were probably introduced by humans, although agreement on this point is not unan- imous. Cannas are characterized by an asymmetric, showy flower with a staminodial labellum, two or three large, relatively unspecialized, petaloid, sterile stami- nodes (in ours), and one partially petaloid fertile stamen. The inferior ovary and capsule are tuberculate, the capsule containing several dark-colored, hard, nearly spherical seeds with stomata on the seed coats and a tangle of emergences on the funiculus. The leaves lack ligules and pulvini. Mucilage canals pass through the rhizomes and unbranched aerial stems. The four genera of Cannaceae recognized by Horaninow were reduced to sections and subgenera of Canna by Petersen (1888) and Baker (1893), re- spectively. In the only revision of the entire genus written during the twentieth century, Kranzlin retained subg. DistEMoN (Bouché) Baker and, subsuming at base, reflexed petals, and 5 dissimilar, petaloid staminodes (labellum to lower lefi, 2 small sterile staminodes to upper right under the letter “‘d’’), x '%; e, frontal view of open flower, labellum to lower right, fertile ae anther i tip of stigma visible in center of flower, x '/; f, detail of flower to show reflexed petals, x '/2; g, expanded staminode, x I: h, fertile stamen with half-anther ne to left margin, < 12; 1, detail of half- Ree after dehiscence, x 3; j, vertical section near center of flower to show petals, staminodes, and style adnate i floral tube. x 1; k, tip of style showing stigmatic line around apex, x 2; 1, diagrammatic cross section of inferior trilocular ovary, 3 septal nectaries visible as black slits near center, the wall finely oe the ovules anat- ropous, axile, x 5; m, mature capsule crowned by persistent sepals, x 2; n, mature seed, x 3: 0, detail of seed coat showing finely pitted surface (with naam not visi- ble), x 25; p, seed in diagrammatic section (cut along line indicated by arrow in “‘n” “‘chalazosperm”’ evenly stippled, x 3; q, seedling still attached to seed, x |. a2 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Baker’s other subgenera in subg. (Eu-)CANNA, divided the latter into sects. BIALATAE and TRIALATAE. Section TRIALATAE (which should be called sect. CANNA) comprised four subsections. Winkler’s infrageneric classification was essentially the same, except that the subsections were designated series. In a revision of the species in northern South America, Segeren & Maas thought it ‘very doubtful” that sections BIALATAE and TRIALATAE should be maintained and did not keep them in their formal treatment, which ts the course accepted in the present paper. Subgenus CANNA (staminodes two or three, in addition to the labellum; labellum reflexed) is represented in the southeastern United States chiefly by Canna flaccida Salisb., 2n = 18, which is probably is se indigenous species. Sometimes called “golden canna,” “yellow canna,’ “bandana of the ever- glades,”’ it occurs on the Coastal Plain from South Carolina to Texas, and in the West Indies, Central America, and northwestern South America. Flowers of this striking species have reflexed petals and very showy, broad, soft, yellow staminodes with undulate margins; the bases of the leaves are gradually tapered. A similar species with a wide range in warm regions south of the United States, C. glauca L.,2n = 18, has been reported from Louisiana and Texas (see Thieret; Godfrey & Tryon 274 [Gu] from South Carolina is probably also referable to this species). Canna glauca (C. angustifolia L., C. stricta Bouché, both syn- onyms from Segeren & Maas) differs from C. flaccida most saliently in its erect petals and narrower staminodes. (Whether or not the staminodes of C. glauca are mottled is a point of disagreement.) Canna indica, indian-shot (with an ample synonymy), 27 = 18, 27 (also probably 36; plus other dubious numbers reported), has escaped cultivation sporadically in the Southeast, as well as in much of tropical Asia, Africa, and elsewhere. This species was cultivated in Europe as early as the sixteenth century. Its capsules and predominantly red flowers marked with (or sometimes entirely?) yellow are smaller than those of the preceding species (e.g., the widths and lengths of the staminodes are about 1-1.5 x 4-6 cm vs. 1.5-2 x 8-10 cm in C. glauca and 8-9 x 9-11 cm in C. flaccida, measurements from Segeren & Maas). Canna indica sometimes pro- duces fertile hybrids with C. glauca despite some meiotic irregularities and diminished fertility. Canna edulis Ker-Gawl., 2n = 18, 27, often cultivated for its edible rhizome, appears in Ward’s checklist of plants native or naturalized in Florida and as a synonym of C. indica in Segeren & Maas. Described by Kriinzlin as “ein uferloses Meer,” the cultivated cannas are an assortment of probably over 1000 horticultural varieties. Most of these fall into two main groups. both of which persist following cultivation or escape in the Southeast: Canna x generalis Bailey does so sporadically in several states, and C. x orchiodes Bailey was reported as growing apart from cultivation in North Carolina by Crutchfield. Flowers of these two diverse, intergrading hy- brid lines are large (reaching 20 cm in diameter in C. x orchiodes) and display a rich array of colors, mostly reddish and yellowish hues, sometimes pastels or white, with the staminodes often spotted, streaked, and bordered. Foliage and stems are bronzed in some. Because C. flaccida is a genetically influential ancestor of C. x orchiodes, confusion between these two 1s especially likely. Canna flaccida has a longer floral tube (over 5 cm vs. ca. 2.5-5 cm in C. x 1984} ROGERS, ZINGIBERALES 33 orchiodes and less than ca. |.5 cm in C. x generalis) and a longer capsule (4- m vs. shorter and sterile in C. x orchiodes). Both C. x orchiodes and C. flaccida differ from C. x generalis and the other species in having petals re- flexed or becoming so and flowers that are generally more delicate and flowing. (For further comparison of the large-flowered cannas, see Bailey, 1923; Crutch- field; and Mukherjee & Khoshoo, 1970d.) Much literature on Canna revolves around elucidating the ancestry and cytology of C. x generalisand C. x orchiodes and morphological changes that have taken place in the course of their development since the mid-1800’s. The history of the cultivars is clouded by loose application of names in early records. From the multifaceted investigations along these lines by Mukherjee and Khoshoo, Canna glauca, C. indica, C. iridiflora Ruiz & Pavon, and C. War- szewiczli A. Dietr. stand out as the most likely principal progenitors of the multiple lineages known as C. x generalis. Selected additional (but not entirely congruous) references useful in this connection are Anonymous (1898), Bailey (1923), and Donahue. Like their four supposed ancestors, most cultivars of Canna x generalis are diploids with normal or nearly normal meiosis. A small proportion are meiot- ically irregular interchange heterozygotes; others are sterile triploids that tend to form trivalents at metaphase I. During the 1890’s, plants of Canna x generalis, in this case ““Crozy cannas,” were crossed with C. flaccida to yield C. x orchiodes. That the genome of C. flaccida is strongly differentiated from those of the putative ancestors of C. x generalis is suggested by the complete sterility of C. x orchiodes, which has highly irregular meiosis in its diploid cultivars and forms few, loose trivalents in the triploids. On the other hand, Kranzlin mentioned fertile hybrids between C. flaccida and C. Warszewiczil. (The accuracy of this report depends upon the accuracy of Kranzlin’s opinion that a plant breeder misapplied the name C. “*nepalensis” to C. flaccida.) Baker (1894) described C. flaccida x iridiflora as the origin of (fertile?) fine garden forms. If one considers the genetic intercompatibility among (and not limited to) the probable ancestors of Canna x generalis, confusion in defining species of Canna is hardly surprising. A related confounding factor is that cannas have long been transported about the world by humans (undoubtedly with operation of the founder principle, release of segregating hybrid progeny into new habitats, vegetative propagation of new forms, and artificial selection). The pantropical complex centered around C. indica is a troublesome example, involving several possible species. A second puzzling example is C. flaccida, which differs only slightly from C. Reevesii described by Lindley from specimens grown from seeds sent to him from China. A further source of taxonomic confusion in Canna is a likely overemphasis of certain characters. For instance, in a series of papers, Honing analyzed the genetics of the coloration of flowers and vegetative parts in C. glauca, C. indica, and other possible species. In 1939 (VI) he explicitly rejected Kranzlin’s clas- sification, concluded that the most obvious differences distinguishing C. indica and certain other “‘bialatae’? depend on only a few Mendelian factors, and proposed placing five names (including C. indica, excluding C. glauca) in syn- 34 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 onymy. (Honing’s studies were summarized by Khoshoo & Mukherjee (1970b), who suggested that members of nearly the same collection of species, including both C. glauca and C. indica, may be differentiated at an ecospecific level.) Red color in the flowers is a monogenic dominant trait influenced in expression by at least 22 other genes and linked to a set with recessive lethal alleles. The presence of two vs. three staminodes in addition to the labellum has repeatedly emerged (e.g., Baker, 1894; Costerus, 19 16a; Jiménez; Kirchoff, 1983b; Segeren & Maas) as a character of questionable taxonomic value, varying within a species and even on a Single plant, yet this character alone distinguishes Kranz- lin’s sections “Bialatae” and “Trialatae.” Species of Canna are predominantly diploids with 2” = 18. Triploidy (which appears to have arisen repeatedly in artificially selected cultivars) and infre- quent tetraploidy have been detected only in species for which diploid counts are also reported. (Compilations of chromosome numbers are available in Segeren & Maas; Mahanty; Khoshoo & Mukherjee, 1970a; Sharma; and Sato). Sato described and illustrated chromosomal morphology for Canna edulis, C. indica, and C. glauca, all 2n = 18, which differ from each other in distributions of lengths of chromosomes, positions of centromeres, and locations of satellites and secondary constrictions. Mahanty concluded from a study of six species that species of Canna form a very homogeneous group. As in Marantaceae, the pollen in Cannaceae is deposited on the style while in the bud, either directly onto or near the stigma. It is widely assumed, although hardly tested, that self-fertilization plays an important role in the breeding system. Mukherjee & Khoshoo (1970b) (see also Darwin) found the few “‘el- emental” species that they studied, cultivated outside of their natural ranges, to be predominantly autogamous, whereas displacement of the anther and stigma necessitated a shift to outbreeding in the cultivated hybrids. (Garden cannas are usually propagated vegetatively.) The fragrant flowers of Canna flaccida open about dusk for a single night throughout the year in South Florida. Swamps, marshes, shores, and pine savannas are the habitats of this species; in the United States C. g/auca and C. indica likewise occupy wet places. This is at least sometimes true of hybrid cultivars growing outside of cultivation. In much of the American tropics and subtropics, including the southeastern United States, Canna is larval host to a skipper butterfly, Ca/podes ethlius (Cramer), which at times is severely destructive to the plants (see Clark & Clark, Cockerell, Evans, Young) and which 1s a pest on the related Maranta arundinacea grown commercially in the West Indies (Purseglove). The larvae, which build tentlike structures protected by rolled or folded leaves, are para- sitized by possibly two genera of wasps. As determined for Canna x generalis and C. indica, the thick wall of the pollen grain has three main layers. Much reduced, the exine, which takes the form of scattered spinules and may also extend around the grain as a thin skin (Hesse & Waha), rests on a thick, bilayered intine. The outer intinous layer, the ‘‘exintine” (Kress & Stone), is honeycombed with radial channels and overlies the unchanneled “endintine.”” Comparable in structure to germinal pores of more typical pollen grains, the “omniaperturate” grains of Canna ies) Gs 1984] ROGERS, ZINGIBERALES appear to have their entire surface suitable for hydration, other interactions with the stigma, and emergence of the pollen tube. The channels in the intine possibly function as reservoirs of compounds connected with incompatibility mechanisms or otherwise with germination of the pollen (Hesse & Waha). For details on the structure of the pollen, see Kress & Stone, Rowley & Skvarla (1974, 1975), Scheer & Franke, Skvarla & Kelly, and Skvarla & Rowley. (Also see discussion of Hedychium in the present paper.) Useful not only as ornamentals, some species of Canna have starchy rhi- zomes, for which especially C. edu/is is used as a source of food for livestock and humans. Possibly first domesticated in northern South America (Gade), this species is cultivated in warm regions around the world, sometimes on a commercial scale. The product from Australia is called “Queensland arrow- root.” A current project of the New York Botanical Garden Institute of Eco- nomic Botany is the development of a clone of Canna with the largest starch grains known from any plant. The hard, more or less spherical, dark-colored seeds have been employed in sundry easily imaginable ways from ammunition to rosary beads, and they have also been used as an ingredient in a substitute for coffee. One example sufficiently underlines their renowned durability: a seed about 600 years old taken from within a rattle recovered from an Argen- tinian tomb grew into a flowering individual of Canna compacta Roscoe (Ler- man & Cigliano). Burning the plants is said to produce an insecticidal smoke. Extracts from C. indica and other species have molluscicidal activity of interest in the control of schistosomes (Mahran et a/.). Additional uses of cannas, especially extensive medicinal applications, are listed in Hegnauer, Kranzlin, Morton (1981), and Perry. REFERENCES: Under ordinal references, see BENTHAM & Hooker, CORRELL & CORRELL, COSTERUS (19 16a, 1916c), Cronauist (1981), Czara, DAGHLIAN, Davis, EICHLER, ERDTMAN, GIBBS, Goprrey & Woorten, HEGNAUER, HEPPER, HORANINOW, HUMPHREY, KIRCHOFF, KNUTH Lone & LAKELA, MAHANTY, Martin, Morton, MULtLer, NEAL, OLATUNJI, PAI (1965a), PERRY, PETERSEN, Bopee: ron et al., PorzTAL, PURSEGLOVE, VW. S. RAo & Donbe, RICK- ETT, Roscoe, SAAD & IBRAHIM, SATO, SCHACHNER, SHARMA, SMALL, TOMLINSON, VENKA- TASUBBAN, WARD, WILLIAMS & HARBORNE, WOODSON & SCHERY, and WUNDERLIN. Anpr®é, E. Italian ae Gard. Chron. III. 18: 703-705. 189 ANONYMOUS. The “Italian” or “orchid-flowering” cannas. aa Chron. II]. 23: 2. 1898. [Includes a aes re ancestry of “‘orchid-flowering” cannas.] Cannas. Missouri Bot. Gard. Bull. 3: 93-99. 1915. ‘Description of 92 cultivars: history and cultivation of cannas.] . Canna. Bull. Natl. Bot. Gard. Lucknow 3: I-15. pls. 1, 2. 1957. [Several varieties described; information on history and cultivation. ] ASHTAKALA, S. S., & R. J. MALONEY. Characterization of anthocyanin pigments in 3 cultivars of garden Canna. Jour. Am. Soc. Hort. Sci. 96: 755-757. 1971.* Baitey, L. H. Morphology of the canna flower. Bot. Gaz. 22: 222, 223. 1896. . Various cultigens, and transfers in nomenclature. Gent. Herb. 1: 113-136. 1923. [Describes Canna « generalis, C. x orchiodes, 118-120; both designated “cultigens nova’’; also includes diagram and photos of flowers.] BAKER, J. G. A synopsis of the species of cannas. Gard. Chron. III. 13: 42, 43, 70, 164, 196, 1893. [Twenty-three species in 4 subgenera. ] 36 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 —. On the species and garden forms of Canna. Jour. Roy. Hort. Soc. 16: 178-188. 1894. [Horticultural and taxonomic history of the genus: reports of several hybrid- izations; discussion of C. indica complex: flowers of Canna compared with those of related families. BArTRAM, W. Travels through North & South Carolina, Georgia, East & West Florida. xxxiv + 522 pp. 8 p/s. Philadelphia. 1791. [C. “/utea” (flaccida?), 155: C. indica, 426.] Betiinc, J. The behavior of homologous chromosomes tn a triploid canna. Proc. Natl. Acad. Sci. U. S. A. 7: 197-201. 1921. [Examined chromosomes from 31 clones of Canna: 25 diploid (3 of these with irregular meiosis), 6 triploid (or nearly so) with different arrangements at metaphase I.] Chromosomes of Canna and of Hemerocallis. Jour. Hered. 16: 465, 466. 1925 [1926]. [Camera lucida drawings of metaphase I in 3 triploid cultivars.] Biacuiy, C. D. A sectorial chimera in the canna. Jour. Hered. 31: 452-454. 1940. [C oncerned with expression of red coloration in foliage and flowers of C. x gene- Boucnue, P. C. Beobachtungen tiber die Gattung ole Linnaea 8: 141-168. 1833 [Genus divided into “Cannae verae” and **Monochila.”’] —. Synoptische Zustammenstellung der Canna- oe so wie zweler davon ge- freuen Gattungen. /bid. 18: 483-495. 1844. [Eurvsty/us, gen. nov.; Distemon, gen. nov.] Crark, A. H., & L. F. Crark. The butterflies of Virginia. Smithson. Misc. Collect. 116(7). Frontisp. + vii + 239 pp. + 30 pls. + map. 1951. [Calpodes ethlius, 183- 185, pl. 23f Includes distributional notes, data on seasonality. and descriptions of infested cannas, oviposition, and the method by which the larva rolls leaves. ] — tL, T. D. A. Notes on the life-history of Ca/podes ethlius. Entomol. News 3: . 1892. eae ICHFIELD, P. J. Taxa collected ne Roanoke Island new to the flora of North Carolina. fc 29: 129-137. 1964. [C. indica, C. x generalis, C. x orchiodes.] Curpeprer, C. W., & H. H. Moon. aie i of the rhizome, stem, ee caf of some horticul 4 sone of Canna in relation to their possible use. U. S. Dep. Agr. Cire 497: 1-21. 1938.* Darwin, C. The effects of cross and self fertilisation in the vegetable kingdom. viii + _ pp. New York. 1877. [Cannaceae., see especially pp. 230-233; studied C. War- zewiczil, this and probably other species of Canna self-fertile. Don: AHUE, J. W. History, oreeae and euler oD of the canna. Am. Hort. Mag. 44: 84-91. 1965. [Includes a chart showing crosses leading to modern cultivated cannas; description and history of cultivars: comments on ae and diseases; instructions for cultivation and artificial pollination. ] Evans, W. H. A catalogue of the American Hesperndae. Part IV. v + 499 pp. pls. 54- SS. London. 1955. [Calpodes ethlius, 402; Evans and some ae authors attribute the name of this butterfly to Stoll rather than to Cramer—it was originally published in a joint work in which the latter probably described the species in question.] Fernatp, M. L. Local plants of the inner Coastal Plain of southeastern Virginia. Part II. Enumeration and discussion of noteworthy species collected. Rhodora 39: 379- 415, 433-459. pls. 474-487. 1937. [C. flaccida, 402: thriving in garden refuse, Isle of Wight County. I have seen Fernald & Long 6172 (cit), the collection that backs up this report, and found 11 to be one of the garden cultivars: see further comments in mi ee D.] Freeman, J. D., AWS. Causey, J. W. SHorRT, & R. R. Haynes. Endangered. threatened. and 7 concern plants of Alabama. 25 pp. Auburn. Alabama. 1979. [C. flaccida, . 10, threatened Furvio, T. E. pi. Recuentos cromosémicos en angiospermas argentinas. IT. Kurtziana 7: 39-42. 1973. [C. compacta, 2n = 18, chromosomes illustrated. } 1984] ROGERS, ZINGIBERALES +) Gape, D. W. Achira, the edible canna, its cultivation and use in the Peruvian Andes. Econ. Bot. 20: 407-415. 1966. [Peruvian remains of C. edulis as old as 2500 B.c.; includes map showing present distribution and hypothetical historical diffusion. } HeENbDERSON, E. Dwarf cannas. Horticulture 44(4): 34, 43. 6. Hesse, M., & M. Wana. The fine structure of the pollen wall in Srrelitzia reginae (Musaceae). Pl. Syst. Evol. 141: 285-298. 1983. [Includes comments on exine in Canna; pollen walls of Strelitzia, Heliconia, Canna, a este ee _ reserva- tions), and certain dicots have thin exines and complex intines having channels in the outer layer (these restricted to | hemisphere in He/iconia and even ce restricted in Tapeinochilos).] Honinc, J. A. Canna crosses. I-VIII. Meded. Landb. Wageningen 26(2). 55 pp. eee 1923:* 32(4). 14 pp. 1928:* 35(1). 26 pp. J pl. rey ** Genetica 15: 23-47. ibid, 21: 70-87, 325-344. 1939; ibid. 23: 277-288. 1943. Howes, F. N. The edible canna (Canna edulis Ker- a ). Bull. Misc. Inf. Kew 1929: 266-268. 1929. [Best 4 in Hawaii 34 tons/acre/year.] Jiménez, R. Cannaceae. . Gomez-Pompa et al., eds., Flora de Veracruz. Fasc. 11. 8 pp. 19 Cc. se © indica. For pollen see Biotica 7: 417-422. 1982.] KHosHoo, T. N. Evolution of garden cannas. Pp. 93-101 in G. Vipa, ea. Symposia Biologica Hungarica. Vol. 12. Evolution in plants. Budapest. 1972. & I. MUKHERJEE. Genetic-evolutionary studies on cultivated cannas. III. Vari- ation in meiotic system. Cellule 68: 109-134. p/s. 1-3. 1970a. [Based mostly on cultivars of C. x generalis, but with information on other species and hybrids; includes description of meiosis at the diploid, triploid, and tetraploid levels.] VI. Origin and evolution of ornamental taxa. Theoret. Appl. Genet. 40: 204-217. 1970b. [Includes diagram outlining history . the two chief groups of garden cannas; in- formation on hybrids outside of the two main cultivated lines; discussion of roles of somatic mutations, polyploidy, en selection, and hybridization in devel- opment of cultiva Kra&nziin, F. Cannaceae. Pflanzenr. IV. 47(Heft 56): 1-77. 1912. [Die ungemeine Variabilitat der Farben in Bliiten, deren Struktur sonst gleich ist, hat, wie immer bei Gartenpflanzen, zu einem Aufstellen von “A rten” gefiihrt, von denen der gréssere Teil vor der Kritik nicht stand halten konnte. . .” (p. 12). Kress, W. J., & D te Srone. Nature of the spuroderai in monocotyledons, with special —_ aa, = oe reference to t ollen grains of Canna and ie Orne 21: 129-148. 1982 [1983]. [C. nde: includes scanning and tr nd light micrographs of pollen gra Lerman, J. C., &E : M. CIGLIANO. New carbon-14 evidence for six hundred years old Canna compacta seed. Nature 232: 568-570. | Linptey, J. Canna Reevesii. Mr. Reeves’s Indian shot. Bot. Register 23: p/. 2004. 1837. [Uncertain that this is distinct from C. flaccida; see also BAKER (1893), KRANZLIN (p. 50).] Lutz, M. L. The gum of Canna. Bot. Gaz. 25: 280, 281. 1898. [Gum forms in thick layers in the interiors of oe MAHRAN, G. G. A. ELt-Ho , M. SAcen, & H. M. Mortawe. Isolation and identification of certain Se eilcscctisl substances in Canna indica L. Jour. Afr. Med. : 107-119. 1977.* [Abstract in Excerpta Bot. 32: 260. 1979. Six isolates tested as RENE activity on the intermediate host of Schistosoma mansoni. Of 2 active substances, | identified as triacontanol; also found were (-sitosterol, stig- masterol, and campesterol.] Ae MouaAmMeb, & H. M. Morawe. Contribution to the molluscicidal activity of Canna species growing in Egypt. Jour. Afr. Med. Pl. 1: 147- 155. 1977.* [Abstract in Excerpta Bot. 32: 259. 1979. C. indica, C. flaccida, and C. orientalis Roscoe.”’] MOookHERJEE, R., & P. N. BHADURI. Cytotaxonomical studies of different varieties of 38 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Canna indica L. Proc. Indian Sci. Congr. 57(3): 289. 1970. [Of 28 cultivars studied, 5 were triploid, the remainder diploid.] Mukuerwee, L., & T. KHosHoo. Genetic-evolutionary studies on cultivated cannas. I. Variation in phenotype. Proc. Indian Natl. Sci. Acad. 36B: 254-270. 1970a. [Studied mostly C. “generalis” (100 cultivars), C. glauca, C. indica, C. tridiflora, C. War- zewiczli, and C. flaccida; these species compared, with chief objective being to trace ae changes in the development of cultivated cannas; “C. generalis” this paper includes all the ornamental cultivars of hybrid origin; distribution map for the entire genus provided.] II. Pollination mechanism and breeding system. /bid. 36B: 271-274. 1970b. IV. Parallelism between natural and induced somatic mu- tations. Radiation Bot. 10: 351-364. 1970c. [C. pallida and C. “generalis’ subjected to “acute gamma radiations’’; spontaneous and induced mutations sometimes result in threadlike staminodes and styles.] V. Intraspecific polyploidy in starch-yielding Canna edulis. Genét. Ibér ee 35-42. cae * VIL. es treatment and hor- ticultural classification. Jour. Bombay Nat. Hist. Soc. 67: —397, 2 lain pls. 1970d. [Includes key to ee. varicties of C. x pen and C. x or- Nair, P. K. K. Pollen grains of cultivated plants. I. Canna Linn. Jour. Indian Bot. Soc. 39: 373-381. 1960. [48 cultivars.] Pollen grains of Indian plants—1. Bull. Natl. Bot. Gard. Lucknow 53: 1-35. 1962. [Canna, 16.] Nikiroroy, Y.L., & G. F. Feoritova. An analysis of pollen of some species and varieties of the genus Canna. (In Russian; English summary.) Bot. Zhur. 67: 166-176. 1982. OFFERUNS, F. J. Meiosis in the pollen mother cells of some cannas. Genetica 18: 1-60. 1936. [C. humilis, C. lutea, C. discolor, C. glauca, and C. glauca x discolor, all 2n = 18 with regular or nearly regular meiosis; includes detailed history of cytological research in Canna, discussion of discrepancies in reported chromosome numbers, and several illustrations of chromosomes; some bivalents show secondary associa- tions.] . aa in the pollen mother cells of Canna glauca “‘pure yellow.” Ibid. 20: 59-65. 1938. Oomen, H. c J. Polyploidy in Canna. Genetica 24: 333-386. 1949. [Diploid and tetraploid C. “aureo-vittata’ and C. “humilis,” and their triploid hybrid (OQomen thought these to be a single species; see HONING (1939 (VI)) for comments on this matter); seed weight a good indicator of ploidy level.] Rowtey, J. R., & J. J. Skvar_a. Origin of - es intine in pollen of Canna. Proc. Electron Microscopy Soc. Am. 32D: 84. 85. 4.[C. x generalis. & The glycocalyx and aes . exine spinules on microspores of Canna. Am. i Bot. 62: 479-485. 1975. ScHeer, U., & W. W. Franke. Annulate lamellae in plant cells: formation during eee and pollen development in Canna generalis Bailey. Planta 107: 145-159. 1972. [Cytoplasm of differentiating pollen grains and pollen mother cells with single or stacked lamellae, these often associated with ribosomes and vesicles and perforated by numerous pore complexes resembling those of the nuclear en- velope; pores ofa stack often aligned, lamellae possibly a degenerative form of rough endoplasmic reticulum; cf. SKVARLA & KELLY SCHUMANN, K, Einige eres zur Nersholoze der Cannabliit Bot. Ges. 6: 55-66. 1888. SEGEREN, W., & P. J. M. Maas. The genus Canna in northern South America. Acta Bot. Neerl. 20: 663-680. 1971. SS B.B., & D. D. Awastui. On the androecium of Canna indica L. Geophytology 5: 30-32. pi. 7. 1975. [Labellum and fertile stamen sole members of the inner whor!.] SRA ARLA, J. J... & A. G. Ketty. Dictyosome development es microsporogenesis in Canna generalis. Pp. 62-67 in J. HesLop-HARRISON, ed.. en: development and physiology. New York. 1971. [Broader end of ee tubular prolifera- — ne. Ber. Deutsch. 1984] ROGERS, ZINGIBERALES 39 tions more extensive than at the smaller gee this end probably contributing vesicles to developing intine; cf. SCHEER & FRANKE.] Row ey. The pollen wall of. Canna and its similarity to the germinal apertures of other pollen. Am. Jour. Bot. 57: 519-529. 1970. [C. x generalis, in- cludes transmission electron micrographs of sectioned grains. ] TuieRet, J. W. Additions to the Louisiana flora. Castanea 36: 219-222. 1971. [C aug C. glauca. THORNE, R. Vascular ae eee unreported from Georgia. Castanea 16: 29- 8. 1951 ne indica, 37, | TomLInson, P. B. The anatomy one Jour. Linn. Soc. Bot. 56: 467-473. 1961. Trivept, B. S., & C. L. VeRMA. Silicified pseudostem of Canna L. from early Eocene “of Deccan Intertrappean beds, M. P., India. Curr. Sci. 39: 442, 443. 1970. [Mentions previous reports of fossil leaf impressions. ] WINKLER, H. Cannaceae. Nat. Pflanzenfam. ed. 2. 15a: 640-654. 1930. Younc, A. M. Notes on the interaction of the skipper butterfly Cal/podes ethlius (Lep- idoptera: Hesperiidae) with its larval host plant Canna edulis (Cannaceae) in Ma- zatlan, State of Sinaloa, Mexico. Jour. N. Y. Entomol. Soc. 90: 99-114. 1982. MARANTACEAE Petersen in Engler & Prantl, Nat. Pflanzenfam. 1888, nom. cons. (ARROWROOT FAMILY) Erect [or scrambling or climbing] small to large herbs of diverse habit. Rhi- zomes branching sympodially, usually with short internodes. Plants glabrous or with usually unicellular hairs, each surrounded at the base by a cluster of inflated epidermal cells. Branches variably developed, sometimes clustered at or above the ground, often widely divergent, each branch bearing (in the fol- lowing order) a basal, 2-keeled prophyll above a short internode, usually a bladeless sheath (interphyll) above a second short internode, a series of petiolate leaves, and a terminal inflorescence often overtopped by axillary growth [in- florescences sometimes arising directly from the rhizome]. Leaves typically distichous, basal or cauline, with open, often auriculate [rarely ligulate] sheaths, and often asymmetric [often variegated] blades, those of leaves on a shoot either homotropous (i.e., the broader halves all on the same side—right or left—of the midrib) [or antitropous, the broader halves on alternating sides]: petiole with a usually pubescent pulvinus at the insertion of the blade; stomata mostly paracytic. Inflorescences highly variable, [completely or] partly tightly condensed to diffusely branched, bracteate; bracts frequently large and colorful, mostly sheathing. Flowers asymmetric, usually in mirror-image pairs, each pair or group of pairs subtended by a bract and a basal prophyll, [with or] without bracteoles. Sepals free. Petals usually unequal and usually connate basally and adnate to the androecium and gynoecium to form a tube. Outer androecial cycle represented by [0 or] 1 or 2 antesepalous, petallike staminodes; the 3 members of the inner cycle all different: the ““cucullate staminode’’* forming a 8A disorderly profusion of names has been applied to the 2 inner sterile staminodes, both of which are hoodlike. Those adopted in the present paper are translated from extensive use in German and have been used in English by other authors. The cucullate staminode (FIG ] g) is also known as the hooded eee Kappenblatt. Kapuzenblatt, or style- bling stamina The callose staminode (Ficure 3, h) has also been called the callosed or callused staminode, Schwielenblatt, and labellu Particularly ee the last term is used for structures in Cannaceae and Zingiberaceae nea are uniquely homologous within each family. 40 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 sterile pouch concave toward the axis of the flower and enclosing the style, with | or 2 appendages inserted on the edge away from the fertile stamen and arching over the concavity; the “callose staminode” bent into a hood, concave toward the axis of the flower (facing the concave side of the cucullate stami- node), usually with a thickening protruding inward from the face of the side nearer the fertile stamen [or with a second thickening on the other side]; the fertile stamen usually borne on the side of the flower adjacent to the other flower of the pair, with a unilocular half-anther inserted laterally on [and sometimes adnate to] a petaloid appendage [this sometimes short and incon- spicuous]. Gynoecium with | [or 3] locules developing and [0 or] | anacam- pylotropous fertile ovule per developing locule, this nearly basal on an axile placenta; nectaries septal; style curled inward upon release from the cucullate staminode, with a specialized pad basal to the stigma for deposition of pollen in the bud; stigma in a depression between 3 variably developed flaps. Fruit indehiscent or dehiscent into 3 valves, dry [or fleshy]. Seeds variable in shape, basally arillate [or not, especially in indehiscent fruits], hard, operculate; en- dosperm formation nuclear, endosperm much reduced or lacking in mature seed, perisperm abundant and penetrated by a curved or straight (sometimes basally forked) canal originating as a vascularized chalazal intrusion into the progressively curving nucellus; embryo U-shaped. Megagametophyte (embryo sac) fundamentally of the Polygonum type, antipodal cells sometimes undergo- ing secondary divisions. Chromosome numbers highly variable, the base num- bers including 4, 6, 9, 11-13. Tyee Genus: Maranta L. A sharply defined family of 25 to 30 genera and probably more than 400 species (usually sorted into two tribes) typical of understory vegetation in wet regions pantropically, but sometimes reaching subtropical or infrequently warm- temperate areas. The greatest concentration of species is in the South American tropics, with lesser centers of distribution in tropical West Africa and tropical Asia. Especially widespread, the genus Thalia L. occurs throughout much of Africa, where it may or may not be indigenous, and from southern South America to the southeastern United States. (The family is also represented in Florida by Maranta arundinacea, an infrequent escape from cultivation.) Out- standing among the genera, which only exceptionally contain more than 20 species, Calathea G. F. W. Meyer probably encompasses considerably more than 130. This genus and TAymocarpus Nicolson, Steyerm., & Sivadasan are the only representatives of the tribe Phrynieae Petersen (ovary with three fertile locules and usually three fertile ovules) in the New World. The tribe Maranteae (ovary with one fertile locule and ovule) is restricted to the New World, except for Thalia, as mentioned above. Marantaceae are unified as a family by their open, usually eligulate leaf sheaths and a pulvinus (or “‘callus’’) at the insertion of the blade. Tomlinson (1961) pointed out that the strongly developed aerial branching system sets the family apart from other Zingiberales (but the presence ofa prophyll followed by one or more incomplete leaves 1s insufficient to distinguish Marantaceae). 1984] ROGERS, ZINGIBERALES 4] The asymmetric flowers, usually borne in homodromous pairs, have a unique explosive pollination mechanism (described below), one or two showy outer staminodes, two hoodlike inner staminodes positioned face to face (one with one or two lateral appendages, the other usually with a thickening protruding from the surface), a half-anther connected to a petaloid appendage (also true of Cannaceae), and a uni- or trilocular ovary with one nearly basal ovule per locule. The abundant nutritive tissue in the usually arillate seed is penetrated by one or two canals. The embryo is curved. While there has been little contention about the delimitation of the Mar- antaceae as a family, the definition and nomenclature of taxa within it have a troubled history. Although fundamental to the studies that followed, Koer- nicke’s monographs (1859, 1862) bear little resemblance in detail to relatively recent treatments. Eichler provided detailed morphological analyses essential for comparing plants as complex as Marantaceae and a firm foundation for his taxonomic treatments of the genera in the New World. These contain very little emphasis on individual species, however. Eichler died while working on the Marantaceae for the Flora Brasiliensis, for which the account of this family was completed by Petersen, who also authored the brief and conservative treatment of the family in the first edition of the Nattirlichen Pflanzenfamilien. Major generic reorganizations, especially among the taxa of the Old World appeared in Schumann’s treatment, which is the most recent revision of the family as a whole. Schumann recognized 26 genera, over twice the number in any of the other works mentioned above. Loesener updated Schumann’s re- vision with a few alterations. In an effort to alleviate persistent confusion in the delimitation of genera, Andersson (198 1a) reevaluated the neotropical gen- era using a wide array of characters and (some taxonomic changes notwith- standing) felt that his studies largely confirmed the validity of Schumann’s genera. Andersson’s (1977, 1981la) proposed view of the early evolution of Mar- antaceae departs markedly from the emphases on a single character and on geographic distributions of the traditional tribal division. According to An- dersson, the family probably originated in Africa (see Holttum for a contrary view); the five informal groups into which he sorted the New World genera hypothetically reflect distinct phyletic lines, each with its closest relatives in the Old World. Tomlinson (1961) failed to distinguish the traditional tribes anatomically, Williams & Harborne found them not to differ in distribution of flavonoid constituents, and Bisson ef a/. could find no cytological support for their maintenance. Marantaceous inflorescences vary from congested capituliform or spicate arrangements to those with clustered subunits or infrequently individual flowers separated by elongated internodes. Spathelike, often colorful bracts are fre- quently arranged in imbricate spirals or distichous rows, or in some species they are widely separated. Growth pattern ranges between strongly monopodial and more or less sympodial. Branching may reach several orders of complexity (with repeated patterns or not) or may be nearly completely suppressed. As is 42 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 evident from two analyses of the architecture of inflorescences in Marantaceae (Eichler; Andersson, 1976), a fundamental uniformity underlies the confusing diversity. Andersson’s attempt to unify the terminology applied to the inflorescence 1s useful if it is borne in mind that the subunits that he recognized are not all constantly discernible throughout the family and that his naming of different levels within the inflorescence deemphasizes repetition in pattern. Like a veg- etative branch, an axis in a marantaceous inflorescence begins with two short internodes, the first bearing a two-keeled prophyll (three-keeled in some flower pairs) and the other bearing an (often-suppressed) interphyll. As the result of repeated condensed basal branching, clusters of branches often falsely appear (e.g., in Thalia dealbata) to be axillary to a single bract. Monopodial axes bearing bracts typically subtending flower pairs or clusters of flower pairs were termed ‘“‘florescences”” by Andersson. The pairs of flowers are borne either singly or in often much-condensed sympodial arrangements (*‘florescence com- ponents’’). Because the morphological nature of the flower pairs remains un- settled, they cannot be counted among taxonomic links to the Cannaceae, which also sometimes have paired flowers. Positions of associated foliar structures, symmetry and time of opening of the flowers, and supposed occasional man- ifestations of the missing terminal flower have been cited as evidence that the flowers of a pair in the Marantaceae are the two lateral members of an ances- trally three-flowered dichasium (see F. Miiller, Eichler). Alternatively, the po- sitions of the bracteoles in some specimens indicate one flower as lateral and the other as terminal (see Andersson, 1976). Kirchoff (1983b) concluded that in Marantaceae the members of a pair originate simply by division of an apical meristem. Arils on seeds of Marantaceae appear to promote dispersal in at least three ways. Schumann (see also Humphrey) noted a correlation between dehiscent fruits and arillate seeds in the family and also stated that expansion of the aril can force apart the valves of such fruits. Eichler observed that the aril jutting rom a partially dehisced fruit can help free it from the enclosing bracts. Bright coloration and deposits of lipids, which are at least sometimes present in marantaceous arils, point to a third role for these structures: the attraction of birds (Ridley) and ants. In a study of species of Ca/lathea and several genera of ants, Horvitz & Beattie found that these insects sometimes (with variation between species) transport the seed by using the aril as a handle. The ants take the seed into their nest (perhaps a favorable. protected microsite for germi- nation), use the aril as food (thus promoting germination), and bury the re- mainder of the seed in a refuse heap near the nest, where the effectively planted seed lies in a bed enriched by decaying organic waste and may be protected by the ants. Prior to the tripping of the pollination mechanism, the style, confined by the cucullate staminode, develops tension, and the half-anther deposits pollen onto a specialized region below the stigma (FiGure 3, k-13). One or two appendages rise from one margin of the cucullate staminode and arch across Its concave side, thereby also extending over the style (FIGURE 3, g). As might be guessed from the appearance of these “triggers” (and as has been confirmed for some — r] ae r] 1984] ROGERS, ZINGIBERALES 43 species), their displacement by a suitable bee releases the springlike style from its confinement. When thus tripped, the style bends suddenly inward toward the callose staminode, which apparently functions in bracing the precisely arranged mechanism, in orienting the insect visitor, and as as anvil for the stroke of the style. The inward-moving stigma scrapes pollen from the insect, which is then at least sometimes smeared with sticky fluid from a glandular region on the style, followed by a dusting with pollen from the pad just behind. According to Kennedy, at least three genera of Marantaceae deposit the pollen on the proboscidial fossa, out of reach of the bee when cleaning itself. After pollination, the curled style and (when present) an appendage beyond the stigma (Ficure 3, j-16) block further loss of nectar and/or autogamy. Euglossine bees. among which are pollinators of many orchids, are the principal pollinators of Marantaceae in the New World (Kennedy) but do not range into the south- eastern United States (Michener). (Supplementary references on pollination are Andersson, 1981a; Eichler; Gris, 1859; Hildebrand; Knuth; H. Miiller; Purse- glove; and Schumann.) The function of the peculiar chalazal canals in the seeds appears to be con- nected with the vascular tissue that defines their course early in development. Because the canals are closed off in the mature seed, they are unlikely to be conduits for air or water (Grootjen). Diverse inclusions of chalazal tissue in seeds of Zingiberaceae. Cannaceae, and Musaceae neither resemble nor initiate such canals. Species of Calathea, especially C. Allouia (Aublet) Lindley, serve as minor sources of starch. Derivation of starch from Maranta arundinacea 1s discussed under that genus. The leaves of species of Calathea yield a wax of high quality (see Ayensu et al.). Several genera, among them Maranta, Calathea (e.g.. C. zebrina (Sims) Lindley, zebra plant), Crenanthe Eichler, and Stromanthe Son- der, are cultivated for their decorative foliage. REFERENCES: Under ordinal references. see Bisson ef al., Costerus (1916c), CRonouist (1981), Czaja, DAGHLIAN, Davis, ERDTMAN, GiBBs, HEGNAUER, HORANINOW, HUMPHREY, KircHorr, KNutH, LonG & LAKELA, MAHANTY, Morton (1976), MULLER, OLATUNII, Par & TILAK, PERRY, PETERSEN, PorzraAL, PURSEGLOVE, A. S. RAo & VERMA, RICKETT. Roscoe, SATO, SCHACHNER, SHARMA, TILAK & Pat, TOMLINSON, VENKATASUBBAN, WIL- LIAMS & HARBORNE, and WUNDERLIN. ANpeERSSON, L. The synflorescence of the Marantaceae. Organization and descriptive terminology. Bot. Not. 129: 39-48. 1976. [Several genera, including Thalia and Maranta, terminology largely adopted from W. Troll. The genus eben (Marantaceae). Op. Bot. 43: 1-113. 1977. [31 species: includes comments on tribes of Marantaceae. The Leen genera of Marantaceae. Circumscription and relationships. Nordic Jour. Bot. 1: 218-245. 198 1a. [Includes tabular comparison of genera, dis- cussion of characters used to distinguish groups, and morphological overview of family. AYENSU, E. S., & R. E. ScHULTES, cochairmen, & AD HOC ADVISORY PANEL. Underex- ploited tropical ae with a cans c value. x + 189 pp. National Acad- emy of Science, Washington, D. C. 6. [Calathea ae possibly other genera as sources of wax, 137-140.] — 44 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Baitey, L. H., E. Z. Battey, & Battey Horrorium Starr. Hortus third. xiv + 1290 pp. New York & London. 1976. [Marantaceae, 713: Maranta, 712, 713. Thalia, 1104.) BatLLon, H. Mémoire sur : symétrie et Porganogénie florale des Marantées. Adansonia 1: 306-327. pl. 11. 1 EICHLe R, A. Beitrage zur Nicaiiene und Systematik der Marantaceen. Abh. Akad. . Berlin Phys. 1883(Abh. 1): 1-99. pls. /-7. 1884. [One of the most important aie on vegetative and reproductive morphology of tomy views on floral morphology in this paper diverge from Eichler’s earlier w Esenpeck, C. G. N. von. Uber die Gattungen Maranta und Thalia. Linnaea 6: 303- 342. pls. 3-5. 1831. Hence on seeds and flowers of 7. dealbata and M. arun- dinacea: descriptions. a illustrations of floral morphology, comparison with each other and with Cann e.] Freire, C. V. Plantas raras. Distribuicdo geographica das Marantaceas. Anais Acad. Brasil. Ci. 6: 197-199. p/. 6. 1934. [Distributions of several genera in Brazil] _ GAGNEPAIN, F. Un cas ie hea d’évolution: la famille des Marantacées. Bull. Soc. Bot. France 77: 190, . 1930. [Characterization of flower, suggestion that fine sixth staminode 1s mee with style. comparison with flowers of Cannaceae, Zin- giberaceae, and other flowers having petaloid androecia. | Grar, A. B. Exot Series 3. ed. 8. 1837 pp. E. Rutherford, New Jersey. 1976. [Mar- antaceae, 1138-1148. and additional scattered references; see also Mfaranta, 1651; Thalia, 1724.] Gris, A. Observations sur la fleur des Marantées. Ann. Sci. Nat. IV. 12: 193-219. pls. 11-14. 1859. [Thalia dealbata, Maranta arundinacea, and species from other gen- era. | Origine et mode de formation des canaux périspermiques dans les Marantces. Bull. Soc. Bot. France 7: 237-239. 1860. [See also GRooTJeN, Gris (under Thalia), and EICHLER. GrooTjJEN, C. J. Development of ovule and seed in Marantaceae. Acta Bot. Neerl. 32: 69-86. 1983. [Species of Thalia (dealbata), Maranta, and 4 extraregional genera; includes literature review and description of development of perisperm canal, seed coat, aril, operculum, and micropylar collar; see also Gris (1860), EicHLer, and SCHACHNER. | HerMANN, W. Die Saag ep der Marantaceen und thre Beziechung zur Tran- soltation: Flora 109: 69-96. 1916. Hitpespranp, F. F. Delpino’s weitere Beobachtungen tiber 7 pene im Pflan- zenreich mit Zusdtzen und Illustrationen. Bot. Zeit. 28: 617-620, p/. 10, figs. 2-9. 1870. [Clear, short, illustrated account of pollination se tn ] Hoittum, R. E. The Marantaceae of Malaya. Gard. Bull. Singapore 13: 254-296. 1951. [Review and criticism of revision by SCHUMANN (p. 263); concerned with taxa in Old World.] Horvirz, C. C. Analysis of how ant behaviors affect germination in a tropical myr- mecochore, Calathea microcephala (P. & E.) Koernicke (Marantaceae): microsite selection and aril removal by neotropical ants, oe Pachycondyla, and Solenopsis (Formicidae). Oecologia 51: 47-52. & . Beattie. Ant dispersal of Calathea ees seeds by carnivorous ponerines (Formicidae) in a tropical rain forest. Am. Jour. Bot. 67: 321-326. 1980. [Two species of Calathea in Mexico.] JonKeR-VeRHOEF, A. M. E., & F. P. Jonker. Notes on the Marantaceae of Suriname Acta Bot. Neerl. 4: 172-182. 1955. ee geniculata compared with 7. trichocalyx: see also Fl. Suriname 1(2): 149-208. 7] KeNNeby, H. Systematics and po iors of Hae closed-flowered” species of Calathea (Marantaceae). Univ. Calif. Publ. Bot. 71: 1-90. p/s. 1-20. 1978. [General remarks 1984] ROGERS, ZINGIBERALES 45 on Marantaceae, including survey of literature on pollination and table summarizing morphological terminology used by other authors; extensive bibliography. ] KircHorr, B. K. Floral development in the Marantaceae: organogenesis. (Abstract.) Bot. Soc. Am. Misc. Ser. Publ. 160: 14. 1981. [See papers by this author under ordinal references. ] Kocu, K. (und Obergiértner LAUCHE). eee Thalia, Phrynium und Calathea. Ber- liner Allgem. Gartenzeit. 25: 161- 857. KOERNICKE, F. Monographiae seucerienn prodromus. Nouv. Mém. Soc. Nat. Moscou 11: 299-362. pls. va 13. 1859. —. Monographiae Marantearum prodromus. Pars altera. Bull. Soc. Nat. Moscou 35: 1-147. 1862. Loesener, T. Marantaceae. Nat. Pflanzenfam. ed. 2. 15a: 654-693. 1930. Matuba, E. Estudio de las plantas de Chiapas vii Marantaceas de Chiapas. Anal. Inst. Biol. Méx. 21: ee 343. 1950. [Calathea, Ischnosiphon, Maranta, Thalia. MicHener, C. D. The social behavior of the bees. A comparative study. xi1 + 404 pp. Cambridge, Meo. 1974. [Chapter 21, The orchid bees (Euglossini), 257- 260. Mosius, M. Merkwiirdige Zeichnungen auf Marantaceenblattern. Ber. Deutsch. Bot. Ges. 36: 263-270, 323-331. pls. 7, 17. 1918. [Species of Calathea: includes illus- trations See patterns. ] Morton, J. Wild plants for survival in South Florida. ed. 4. 80 pp. Miami. 1977 Ivana arundinacea, 22: Thalia geniculata, 25 (rootstock of young plants boiled and eaten). MULLER, . Die Bliithenpaare der Marantaceen. Ber. Deutsch. Bot. Ges. 3: 54-56. 1885. [Morphological support for EicHLer’s belief that the flowers ofa pair are both lateral. ] Nicotson, D. H., J. A. STEYERMARK, & M. SIVADASAN. Thymocarpus cannoides (Mar- antaceae), a new genus and species from Venezuela and Brazil. Brittonia 33: 22-24. 1981. a H.N. The dispersal of plants throughout the world. Frontisp. + xx + 744 pp. . 1-22. Ashford, Kent (England). 1930. [M ras 424, 425)] ScHU ee K. Marantaceae. Pflanzenr. 1V. 48(Heft 11): 1-184. 1902. Tomuinson, P. B. Morphological and anatomical ee of the Marantaceae. Jour. Linn. Soc. Bot. 58: 55-78. 1961. [Includes diagramed analysis of vegetative branching system and brief discussion (with references) of the role of the pulvinus; see also TOMLINSON (1969, under ordinal references). ] TRAN VAN Nam. Sur le callus des Marantacées. (English summary.) Bull. Soc. Bot. France 121: 97-108. 1974. [Includes review of literature concerned with anatomy of the pulvinus, a developmental study on the pulvinus of several species (none ours), and anatomical illustrations. ] |. Thalia Linnaeus, Sp. Pl. 2: 1193. 1753: Gen. Pl. ed. 5. 3. 1754. Usually tall (1-4 m in the southeastern United States), erect, large-leaved herbs of wet habitats. Glabrous or with soft hairs on axes of the inflorescence. bracts, floral parts, fruits, and leaves. Foliage leaves mostly or entirely basal, petiolate, homotropous. Inflorescences variably and usually richly branched, often with 3 or more axes falsely appearing to arise from a single node, the main axis clearly monopodial (7. dea/bata), or forming pseudodichotomies with axillary branches (7. geniculata), the lateral branches with further bifur- cations or not, the penultimate units conspicuous kinked or zigzag axes (“‘flo- rescences”’ or “rachises’’) bearing a pair of flowers at each kink. Flowers of 46 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Figure 3. Thalia.a—m, 7. genicu/ata: a, portion of inflorescence, many flowers ree other flower buds enclosed within bracts (spathe and prophyll), cf. Figure 4, d, x '2; b, pair of tripped flowers (mirror images) enclosed within bracts (1 and 2), the ona ae axis of inflorescence branch (unshaded circle) removed just above base of bracts, the 1984] ROGERS, ZINGIBERALES 47 each pair laterally appressed and enclosed in a pair of unequal bracts, the smaller bract adjacent to the rachis (this bract the prophyll on the condensed shoot bearing the flower pair), the condensed shoot subtended by the larger, abaxial, boat-shaped bract (spathe), the spathes of several pairs often imbricate along the rachis while immature, spathes and prophylls both falling off with the fruits, leaving the rachis naked; other foliar organs in the inflorescence caducous or persistent. Corolla and androecium purplish or blue, sometimes pink [or white]. Sepals very small, distinct, hyaline. Corolla lobes distinct or nearly so, nearly equal to unequal, variably fused at the base with the androe- cium. Outer staminode constricted basally, showy and protruding, borne nearly abaxially to the rachis; cucullate staminode with 2 appendages, markedly con- nate with the callose staminode, this a very broad, bilobed hood provided at the edge adjacent to the fertile stamen (with which it is sometimes strongly connate) with an oblique, inward-jutting thickening; fertile stamen with its half-anther inserted laterally on a variably developed petaloid lobe. Gynoecium unilocular by abortion, with | fertile ovule; style twisted when triggered, free from the androecium, with an appendage derived from the initially adaxial rim of the stigmatic cleft projecting back into the flower after pollination. Fruit ellipsoid to subglobose, indehiscent, with a thin pericarp. Seed single, subglo- bose or ellipsoid, sometimes slightly flattened, with a smallish aril, the peri- sperm penetrated by 2 curved canals originating from a basal bifurcation and flanking the embryo. Type species: 7. geniculata L. (Name commemorating Johann Thal, ca. 1542-1583(?), German physician and naturalist, who died after falling from a carriage.) A cohesive, predominantly South American genus of seven or a few more species ranging from the southern United States, where two species are indig- enous, to the West Indies, Argentina, and Uruguay, and represented throughout much of tropical (especially western) Africa by the often-mentioned “Thalia Welwitschii Ridley” and two other possible species (see Hepper, Koechlin, Gagnepain). As discussed below, it is likely that the widespread African pop- ulations resulted from human activity. Said to be the largest-leaved herb native to the southeastern United States, floral le numbered (see below for key to numbers), x 5: c, right-hand flower from “bx 5: d-k, parts of flower from ‘‘c,” x 3; 1, diagrammatic cross section of ovary, Sire as in flower (c), the fertile ovule above with its micropyle toward the center septal nectary, the 3 septal nectaries and 2 empty, compressed locules below, x 12 fertile, basal ovule, the micropyle to lower left, 2: Key to floral parts in “b’-"k”: 1, outer, lower bract (spathe) enclosing pair of flowers; 2, upper, inner bract (prophyll) | g pair of flowers; 3, upper, adaxial petal; 4, lateral petal (second lateral petal not visible): 5, outer staminode; 6-10, stamens of inner whorl— 6, cucullate staminode (encloses style until pollination mechanism 1s triggered); 7, pet- aloid lobe, which together with callused lobe, 8, forms callose staminode, this connate with 9, fertile half-stamen, and 10, sterile apace (the callose staminode connate at the other margin with 6); 11, lateral sepal; 12, inferior ovary; 13, pollen-bearing area of e (in ‘‘b” with pollen); 14, stigmatic aa after pollination (note pollen); 15, style; 16, stylar appendage 48 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Figure 4. Thalia. a-1, 7. geniculata: a, young leafy shoot (small)—note bladeless sheathing leaf at base, overlapping sheaths, and pulvinus between blade and petiole. x “4: b, diagrammatic cross section of sheaths of 2 leaves and ofa third, much younger leaf to show vertical plates of tissue (vascular bundles omitted) plus horizontal dia- phragms (stippled), x 1'4: c, detail of a diaphragm from “‘b” to show connecting stellate cells with openings between, x 25; d, tip of inflorescence branch with 2 mature fruits, each enclosed by pointed outer and frayed inner bracts (spathe and prophyll, respec- tively), x 2: e, mature fruit with marcescent floral parts on top, x 2: f. seed, removed from fruit—note aril at base, x 2: g, base of seed to show aril, x 2; h, diagrammatic vertical section of seed (through arrow 1n “‘1”’), thick part of aril to left at base, invagination of chalaza in black, perisperm stippled, curved embryo unshaded, outer seed coat heavy black line, inner seed coat unshaded, x 3:1, cross section of seed (oriented as in “*h’’) at level of arrow in “h,” showing seed coats, perisperm, embryo, and chalazal canals in perisperm, x 3. j-o, 7. dea/hata: j, inflorescence axes after fruits have fallen to show 1984] ROGERS, ZINGIBERALES 49 Thalia geniculata L. (T. divaricata Chapman, T. trichocalyx Gagnep., 7. Wel- witschil) in this country is restricted to Florida but extends southward from Mexico and the West Indies approximately to Buenos Aires, Argentina. It is essentially absent from the drainages of the Amazon and upper Orinoco rivers. A distribution map of this species broadly defined with superposed highly variable but poorly correlated characters 1s provided in Andersson’s revision of the “Thalia geniculata complex.” The main problem tackled in this study was a taxonomic evaluation of the three African species of Thalia, the mor- phological divergence of which Andersson attributed to a ‘bottle neck effect” following supposed introduction of 7. genicu/ata into Africa, possibly in the ballast of ships sailing from the Greater Antilles during the slave trade. As a consequence of this hypothetically recent and artificial separation, all African thalias were synonymized with 7. geniculata. Andersson mentioned that plants fitting the original description of the tropical American 7. trichocalyx grow in Florida, but he also included this species under 7. geniculata. Thalia geniculata f. rheumoides Shuey, with red coloration on the leaves, inflorescences, an particularly the petioles, occurs in east-central Florida. Miége’s report of 2” = 18 for 7. geniculata is based on material from Senegal; the provenance of the cue ae by Mahanty for a photograph of the chromosomes (2 = 26) is not spec pats oe hae Fraser ex Roscoe? (7. barbata Small), powdery thalia, 2” = 12, grows on the Coastal Plain from South Carolina to Texas, in Oklahoma, Arkansas, and Missouri. This species differs most saliently from 7. geniculata in having more compact, more rigid inflorescences with a less dichotomous aspect (internodes on the rachis 2-3 mm vs. 5-16 mm); smaller flowers and bracts (the spathe of each pair mostly less than 1.5 cm long vs. mostly longer); and the pruinose bloom for which it 1s named. Delimitation of Thalia, which has no obviously close relatives, has been less troublesome than that of other genera of Marantaceae. Thalias are usually large herbs of wet places; they have homotropous leaves and paniclelike inflores- cences with distinctive zigzag axes. The purplish flowers (ours) are borne in tight, bract-enclosed pairs without bracteoles; each has one conspicuous outer staminode, a very short corolla tube, minute sepals, a cucullate staminode with °T have not seen an illustration of this species said by Roscoe to have been published by Fraser in Ic. Bot. Gard. Liverpool, probably in 1794. branching pattern —compare with ‘‘d” and with Ficure 3, a, x 2; k, tip of inflorescence xis with a single fruit enclosed at base by prophyll and subtending spathe and tipped _ marcescent floral parts, x 2; 1, side view of seed, showing basal aril, x 2; m, base of eed to show aril, x 2; n, vertical section of seed (at level marked by arrow in “‘o”’), aoa thick part - aril to left, outer seed coat wavy, inner seed coat unshaded, basal invagination of chalaza in black, perisperm stippled, embryo unshaded (note that the 2 chalazal canals in perisperm branch from the inv aeination Ne here), x 3; 0, Cross section of seed (at level marked in “‘n’’), showing curved emb nshaded) in perisperm with parallel chalazal canals (much longer than in “1 and thus each cut twice) in black, x 3 50 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 two (vs. usually one in other Marantaceae) appendages, a style with a large backwardly pointing appendage, and a unilocular ovary. The seed has a usually small aril and two (vs. usually one) curved chalazal canals. Andersson (198 La), confirmed the taxonomic isolation of 7halia and compared it with his ““Myr- osma group” of genera, Halopegia K. Schum., and Hypselodelphys (K. Schum.) Milne-Redhead, from all of which Thalia is distinguishable by the character- ization above. In the principal revision of Thalia, Schumann named four subgenera, which were retained by Loesener: EUTHALIA (containing our 7. dealbata), ARTHRO- THALIA (containing only 7. geniculata), SAROTHALIA, and ANOMOTHALIA. In view of the small size of the genus (and because the first two subgeneric names contravene the ICBN, 1978) their continued recognition is of questionable value. The two species of Thalia in the Southeast grow in or bordering ponds, streams, swamps, ditches, hammocks, and other wet places. Both flower from spring through autumn. Andersson (1981b), on the basis of observations in the field, thought plants of 7. geniculata to be self compatible and geitono- gamous. In the Southeast various insects reportedly visit flowers of Thalia; however, only carpenter bees (Xyv/ocopa) are known to trigger the pollination mechanism (D. Austin, see acknowledgments). A gas-filled space between the seed and pericarp in 7. dea/bata, the canals in the perisperm, and the persistent waxy bracts may promote dispersal by water (see Grootjen Seeds of Thalia “divaricata” (geniculata) are ie eaten by ducks in Florida (McAtee). The inner portion of the boiled rootstock of 7. geniculata is said to be suitable as food for humans (Morton, 1977). Thalia dealbata is sometimes cultivated in aquatic gardens. REFERENCES: Under ordinal references, see CoRRELL & CORRELL, GODFREY & Wooten, HEPPER, Humpurey, MAHANTY. MARTIN, SAAD & IBRAHIM, and SCHACHNER; also see ordinal references listed at beginning of family references; under family references, see ANDERSSON 8 la), EICHLER, ESENBECK, GROOTJEN, KOERNICKE, LOFSENER, MORTON, SCHUMANN, and TOMLINSON. /~ an L. Revision of the Thalia geniculata complex (Marantaceae). Nordic Jour. : 48-56. 1981b. [Entire complex made up of a single polymorphic species no ee subdivision of which is justified: 7. geniculata redescribed, full synonymy provided; includes distribution map with superposed distribution of characters for New World, scatter diagram, histograms, table of allegedly distinctive characters in some synonymous taxa, photographs of plants, illustrations of flowers and seeds, and typification.] ~~ J. J. The birds of America. xii pp. + 500 pls. + pp. xui-xxvi. New York. 7. [T. dealbata, pl. 183, : xvil; colored plate. ] ae o G. Scitamineae. Jn; W. T. THiseLton-Dyer, ed., Trop. Afr. 7: 293-331. 1898. [7. coerulea, T. Vu ehuiiscnie T. geniculata, 313 CHAPMAN, A. W. Flora of the southern United States. ed. 3. XXXIX + 655 oo Massachusetts. 1897. [7. dealbata, T. divaricata, in Cannaceae, 491, 492.] Dean, B. E., A. Mason, & J. L. THomas. Wildflowers of Alabama and aint states. xxii + 230 pp. University. Alabama. 1973. [7. dealbata, 32, 33, color photo 1984] ROGERS, ZINGIBERALES 5 GaGnepain, M. F. Zingibéracées et Marantacées nouvelles de lherbier du muséum (1 2° note). Bull. Soc. Bot. France 51: 164-182. 1904. [7. dipetala, sp. nov., 181, compared with 7. geniculata, cf. ANDERSSON, 1981b.] Gris, A. Origine des cannaux périspermiques dans le Thalia dealbata. Bull. Soc. Bot France 7: 875. 1860. [See also Gris (1860) and Grootien (both under family ref erences). ] KoecHLIn, J. Marantacées. Jn; A. AuBREVvILLE. Fl. Gabon 9: 91-158. 1964. [Thalia, 139, 140.] McAtee, W. L. Eleven important wild-duck foods. Bull. U. S$. Dep. Agr. 205: 1-25 1915. [Thalia, 7-9. Mice, J. Troisiéme liste de nombres chromosomiques d’espéces d’Afrique occidentale. Ann. | Sci. Univ. Dakar 5: 75-85. 1960. [7. geniculata, 81; chromosomes illus- oo trated, 8 To T. ale dealbata. Florist Pomol. 1872: 228-230. 1872. [Drawing, short de- scription, information on cultivation SHuuey, A. G. Ared-petioled form of Thelia geniculata L. from central Florida. Rhodora 77: 210-212. 1975. [Brevard and Palm Beach counties Sims, J. Thalia dealbata. Mealy thalia. Curtis’s Bot. Mag. Al: pl. 1690. 1815 (1814°). 2. Maranta Linnaeus, Sp. Pl. 1: 2. 1753; Gen. Pl. ed. 5. 2. Wels Perennial, or perhaps sometimes annual, erect [or prostrate] herbs of diverse habit, [some species without aerial stems or branched only basally,] when branched aerially the main shoot and axillary growth often divaricate-pseu- dodichotomous, usually terminating in inflorescences. Rhizome fleshy, with scale-covered stolons (M. arundinacea). [Glabrous or] with pale indument of variable length, texture, and density on stems, leaves, and ovary (MW. arundi- nacea). Leaves homotropous [except in \/. Ruiziana Koern.], separated by distinct internodes, variegated in some species. basal and cauline [or entirely basal]. extremely diverse in shape (lanceolate to elliptic with the apex acuminate in MW. arundinacea), the sheath sometimes extending to the pulvinus, this usually with an adaxial tuft of hairs. Inflorescences racemiform to diffusely branched, with the axillary and main shoots divergent (in the uncrowded 1n- florescences of M. arundinacea and some other species, the pattern of growth neither clearly monopodial nor clearly sympodial) [some species with compact. spicate branches (“‘florescences’’), these actinomorphic or bisymmetrical]: flow- er pairs borne singly or in few-membered clusters (““florescence components”), each pair with a prophyll but no apparent interphyll, and no [or rudimentary] bracteoles, the pedicels of the 2 flowers and their common stalk of variable lengths, | flower apparently terminal and on a longer pedicel, the other ap- parently lateral and on a shorter pedicel; bracts sheathing, persistent or ca- ducous. Flowers white [to bluish, purplish, or otherwise]. Sepals free, equal, acute, fibrous. Corolla with a usually long tube, this usually gibbous at the base and bent and adnate to the androecium and style; corolla lobes subequal, cucullate at the apex. Outer staminodes 2 (or 3), unequal, conspicuous and flaglike (obovate and usually emarginate in W. arundinacea): cucullate stam- inode with | lateral appendage: callose staminode often 2-lobed; fertile stamen with the petaloid half basally connate with and apically wrapped around the cucullate staminode (4. arundinacea); pollen grains inaperturate or possibly JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 ‘nn bho sometimes uniaperturate. Ovary |-locular by abortion with | fertile ovule; stigma in a usually funnel-shaped hollow formed by a pair of lips at the end of the style. Fruit indehiscent or sometimes 3-valved, ellipsoid, 3-angled in transverse aspect, with | seed (some cultivars of W/. arundinacea seedless). Seed rectangular in longitudinal view, triangular in transverse view, with a pale yellow aril; perisperm canal straight, bifurcate apically. Type species: MW. arun- dinacea L. (the only species included by Linnaeus in the Species Plantarum). (Named for the botanist Bartolommeo Maranta, died ca. 1560, a native of Venosa, Italy.) About 20 species, many of them poorly defined, in four questionable sub- genera, indigenous to tropical America, especially Brazil, with Maranta Fried- richsthaliana Koern. endemic to Costa Rica and VW. arundinacea and M. di- varicata Roscoe (= M. arundinacea?) also in Central America. No species 1s clearly native to the West Indies (see Howard for comments on three species collected in the Lesser Antilles). Since Vf. arundinacea has escaped cultivation and become naturalized sporadically in warm portions of both the Old and New Worlds, its original distribution can only tentatively and vaguely be stated as tropical South America and possibly Central America. Whether or not this species preceded Europeans in the West Indies is uncertain; 1t was clearly under cultivation there in the 1600's (cf. Hodge & Taylor, Purseglove, Sturtevant). Cultivated at least as far north as northern Florida, \/. arundinacea has evi- dently become a minor element in the flora of South Florida. Small (p. 361) described it as occurring on “hammocks and moist soil, S. pen. Fla..”” and in a few more recent publications (Ward; Morton, 1977). this species 1s accepted as growing without cultivation in Florida. However, its meager representation in other checklists and consulted sources should be noted. This representative ofsubg. MARANTA (divaricately branched: bracts few. not imbricated, convolute around the axis; flower pairs on long pedicels) has uncrowded, highly branched growth, leaves with long-attenuate apices, white flowers with the ovaries gla- brous or pubescent on the corners, and thickened stolons. 1859 and 1862 Koernicke envisioned \/aranta as a broadly circumscribed genus made up of four “Untergattungen” or (in his second paper) sections: EUMARANTA, STROMANTHE (Sonder) Koern., SARANTHE Regel & Koern., and XorOLepPIS Koern. Eichler retained part of Koernicke’s sect. EUMARANTA as the genus Maranta and sorted the remainder of the former genus, with changes in the groupings, chiefly into the genera Stromanthe Sonder, Saranthe (Regel & Koern.) Eichler, and Ctenanthe Eichler, all of which have been maintained as genera aga alana eran in delimitation in subsequent revisions. Using characte ith the form and arrangement of bracts, coloration of leaves, habit, and structure of the inflorescence. Schumann named four new subgenera under Maranta: AUTOMARANTA (1.¢€., MARANTA), KOERNICKEA, CAL- ATHEASTRUM, and FRIEDRICHSTHALI Andersson (1981la), who re ered Sehtimana’ s subgenera, placed Maranta alongside Monophyllanthe K. Schum. to form his informal “.Waranta group” and indicated that the two genera perhaps ought to be merged. Species of the wa) Ww 1984] ROGERS, ZINGIBERALES Maranta group tend to have long floral tubes; fibrous, acute sepals: the cucullate staminode with only one appendage: a funnel-shaped stigmatic orifice; flower pairs with elongate axes and without interphylls apis (usually) bracteoles; an apically bifurcate perisperm canal in the solitary, mature seed; and usually homotropous leaves. Maranta differs from Wea according to An- dersson, in having two equally developed outer staminodes (vs. one staminode reduced or absent) and in details of habit. Andersson concluded that the closest affinities of the Maranta group are with the African genus Marantochloa Brongn. & Gris, which differs in having three fertile ovules, instead of only one. Saranthe, Ctenanthe, and Stromanthe, placed in the **Myrosma group” by Andersson, are evidently not strongly differentiated either from each other or from the Maranta group. With exceptions, these genera differ from Maranta in having proportionately shorter floral tubes, usually shorter internodes and more frequent bracteoles in the flower pairs, and a tendency to have the bracts on branches of the inflorescence (florescences) in a mMonosymmetric arrange- ment (vs. radially or bilaterally symmetrical). Crenanthe and Stromanthe differ further from most species of Maranta in having antitropous leaves. Any generalization about cytology in Maranta is hampered by reports on dubiously identified specimens, even at the generic level, and by substantial . heterogeneity. Vouchered survey work is needed. From the infor- we have, it appears that somatic chromosome numbers range between 18 and 52. (A diploid number of 8 was reported for M. “nitida-picta’” —see Venkatasubban.) As judged from very limited data, cytological differences be- tween species of Maranta extend to the lengths of chromosomes and the po- sitions of centromeres. In VW. arundinacea, 2n = 18 (46, 48, 52 also reported). Satd encountered two long pairs of chromosomes with submedian centromeres, a medium-long pair with median centromeres, five short pairs (one with sub- median centromeres, the remaining four with subterminal), and one very short pair with terminal centromeres. According to one account in the collection of correspondence concerning Maranta arundinacea (Anonymous, 1893), t = name ‘“arrowroot” may have originated from the Carib ara-riuta, “mealy root’: alternatively, the name may stem from use of the rhizome in treating wounds from poisoned arrows. ““Ar- rowroot” has become generalized to other plants with starch-yielding rhizomes, such as Canna edulis. Produced in tropical countries worldwide, chiefly on the West Indian island of St. Vincent, true arrowroot starch is extracted from the tough, fibrous rhi- zomes of Maranta arundinacea. Easily digestible, the starch is particularly ani for feeding infants and persons with special dietetic requirements. Additionally, it is or has been used like other starches, such as for starching laundry, in glue, and in the preparation of powders for the skin. Among the many medicinal uses are application internally and externally to counteract poisons and in poultices to treat various dermatological afflictions. Species of Maranta—among them M. bicolor Ker-Gawl. and the prayer plants, M. /eu- coneura Morren (including var. ervthroneura Bunting)—are popular in homes and greenhouses for their decorative, sometimes variegated foliage. At least 54 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 one cultivar of VW. arundinacea has variegated leaves. (Additional references concerned with uses for Maranta are Ayensu, Hamilton, Hodge & Taylor. Morton (1977), Neal. Purseglove. and Sturtevant.) REFERENCES: Under ordinal references, see MORTON, NEAL, PuRSEGLOVE. SAAD & IBRAHIM, SATO, SMALL, STANDLEY & STEYERMARK, VENKATASUBBAN, and Warp; also see ordinal refer- ences listed at beginning of family references: under family references. see ANDERSSON (198la), Battery et al, ErcHtrr, ESpnpeck, Korrnicke, Morton, SCHUMANN. and TOMLINSON. Anonymous. St. Vincent arrowroot. Bull. Misc. Inf. Kew 1893: 191-204. 1893. [Col- lection of correspondence. See also pp. 331-333: cultivation, processing, yield, uses for starch, soils, diseases. | Calathea & Maranta. Bull. Natl. Bot. Gard. Lucknow 33: 1-10. 4 unnumbered ls. 1959. [Includes descriptions of cultivated marantas, instructions for cultivation, comments on pests and diseases, drawings of leave Ayensul, E. 8. Medicinal plants of the West Indies. 282 pp. Algonac, Michigan. 1981. [Varanta, 122: used for digestive disorders. sunburns. poisoning, irritation from manchineel, and other ailments. | DemutH, E. Die Blattbewegungen der ees leuconeura Morr. var. Kerchoveana Morr. Phyton Buenos Aires 9: 117-135. 195 Denmark, H. A., & E. NICKERSON. A tal eer mite, Steneotarsonemus furcatus De Leon, a serious pest on Maranta sp. and Calathea sp. (Acarina: Tarsonemidae). Proc. Florida State Hort. Soc. 94: 70-72. 1981 [1982]. [Concerned with cultivated plants; includes description and photographs of damage: efficacy of various pesticides tested. ] Haminron, W. On the Maranta arundinacea, and its application as an antidote to animal and vegetable poisons. Pharm. Jour. II]. 6: 25-28. 1847. [Starch vield com- pared with that of other plants. Hopar, W. H. & D. Taytor. The ethnobotany of the Island Caribs of Dominica. Webbia 12: 513-644. 1957. (Af. arundinacea, 550-552. magical and medicinal uses. comments on varieties, role of arrowroot in rites.] eam Hoim, T. Sciaphilous plant- ee Beth. Bot. Centralbl. —89. pls. 1-3. 1927. [M. arundinacea, 43, 44, pl. 1, fig. 4: description cn of leaf anatomy. ] Howarp, R. A. Flora of the Lesser Antilles. Vol. + 586 pp. Jamaica Plain, Mas- — sachuseuts. 1979, [Maranta, 548-550: M. ae. M. gibba, M. Ruiziana (cul- ed)—the last 2 species only known (poorly) in the Lesser Antilles from old collections } Pray, T. R. Marginal growth of leaves in monocotyledons: Hosta, Maranta and Phil- eae Phytomorphology 7: aaa 1957 [1958]. LVF. bicolor. superficial initials of — meristems on leaves rarely, if ever, contribute to internal tissues of leaf] SHARMA, A. K., & N. K. BHATTACHARYYA, ‘In ea in chromosome oa in species on: lis and Calathea. Proc. Natt t. Sct. India 24B: 101-117. pl. &. 1958. [Four species of *Maranta,” 3 of which pane to be aiamiber: of € ae includes a or of chromosome numbers from other literature. ] Sims, J. Maranta arundinacea. Indian arrow-root. Curtis’s Bot. Mag. 49: p/. 2307. 1822 [Colored plate shows foliage, branching. inflorescence. and floral dissection. ] SINGH, K. Use of arrow-root powder in starch gel electrophoresis. see 34: 551 1978. [Arrowroot starch used for preparation of gels. cheaper and in some ways better than potato starch; note, however, botanical source of starch not specified. ] STURTEVANT, W. C. History and ethnography of some West Indian starches. Pp. - 199 in P. J. UEKO & G. W. Dimpirpsy, eds., The domestication and exploitation of > nn Loa) 1984] ROGERS, ZINGIBERALES plants as animals. Chicago. 1969. [M/. arundinacea, 184-189; discusses origin of common name, history of the plant in the West Indies, a. uses, processing: malice a worldwide list of localities where arrowroot 1s grow ARNOLD cS eae 22 Divinity AVENUE CAMBRIDGE, eee HUSETTS 02138 ADDENDUM After this paper was completed. a specimen of Thalia geniculata cultivated indoors at the Arnold Arboretum flowered for the first time, allowing us to make the following observations. In bud, the anther deposits large (ca. 100 um in diameter when fresh) pollen grains onto the style within the cucullate staminode. Possibly functioning as a trigger, the stylar appendage in the open untripped flower rises as a steep uphill floor at the entrance of the floral tube. The pollination mechanism is extremely sensitive and is easily sprung by various minor disturbances. After being (artificially) triggered, the style twists and comes to rest blocking the floral tube, with the pollen-bearing portion against the outside of a pouch formed by a membrane extending between the callose staminode and the thickened lobe on it. Presence of numerous pollen grains in the stigmatic cleft suggests that self-pollination can occur in the absence of a pollinator during the stroke of the style. Consistent with this possibility, artificially triggered flowers readily formed fruit when bagged in bud or immediately following triggering. (It was impossible to bag flowers singly.) I have not ruled out the presence of pollen in the stigmatic cleft prior to release of the style, other forms of autogamy, or apomixis. YING ET AL., DIPHYLLEIA a7 A MONOGRAPH OF DIPHYLLEIA (BERBERIDACEAE) TsSUN-SHEN YING, SUSUMU TERABAYASHI, AND DAviID E. BOUFFORD DipHYLLeIA Michaux is one of approximately 120 genera with members exhibiting the classic eastern Asian—eastern North American disjunction pat- tern. It was one of the first examples of this pattern that Asa Gray (1859) pointed out when he listed D. grayi F. Schmidt (as D. cymosa Michaux) as one of a number of taxa restricted to these two widely separated areas. Three species of Diphylleia (D. cymosa, D. grayi, and D. sinensis H. L. Li) are pres- ently recognized, each occurring in separate, disjunct regions. Despite this well- known distribution pattern and the interest it has generated, there have been relatively few detailed studies of the taxa involved. The following account is an attempt to bring together the scattered information on Diphylleia and to point out areas where further studies might result in a better understanding of this disjunction pattern. Floristically, eastern Asia and eastern North America are two of the largest and most complex relictual areas for the once-widespread Mixed Mesophytic Forests that circled the globe at middle and high latitudes in the Northern Hemisphere during the Tertiary (Wood, 1972). These forests and their floras become extinct in large intervening areas during the orogenies and climatic deterioration of the late Tertiary and the glaciations of the Pleistocene. Di- Ww fossil histories (e.g., Fagus L., Liriodendron L., Magnolia L., Cercidiphyllum Sieb. & Zucc., Cornus L. subg. Thelycrania Endl., Tsuga Carr.) to have been a member of these forests, even though the components of the present forest may have had very different histories. Although it might be argued that the current distribution of Diphyileia is the result of long-distance dispersal, it seems more likely that the berry fruits enabled a more continuous and wide- spread distribution during the Tertiary by dispersal over shorter distances. The present localized distribution of D. cvmosa in the southern Appalachian Moun- tains of the southeastern United States is unexplainable. In cultivation plants of this species are able to grow far north of their present range. It may be that they are unable to compete in nature with other, better-adapted plants farther north, or that they have been unable to bridge intervening unsuitable habitats since the melting of the last continental glaciers to reach the seemingly suitable habitats occurring outside of their present range. In addition to Diphylleia, other genera and species of Berberidaceae s./. (Caulophyllum Michaux, Jeffersonia Bartram, Podophyllum L., and close rel- atives of Berberis canadensis Miller) are well known for their discontinuous ) President and Fellows of Harvard College, 1984. ae of the Arnold Arboretum 65: 57-94. sree 1984. 58 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 distribution in eastern North America and eastern Asia. With the exception of Berberis, these genera are all perennial, rhizomatous herbs that grow in rich, moist soils in mixed deciduous forests. ECOLOGY The three species of Diphyileia occupy similar habitats in the eastern United States, Japan, and China. All are found on gentle to steep slopes in seepages or along streams. They grow in brown forest soils or podzols that are rich in organic matter and characteristically acidic (pH 4.5-7.0 for D. sinensis in the Shennongjia Forest District in central China (Shennongjia Bot. Exped. 10276 and 25107); 4.0-5.8 for = cymosa in samples from Watauga County, North Carolina (Boufford & E. W. Wood 16310), and Rabun County. Georgia (Bouf- ford & E. W. Wood oe 3). in the United States). The plants are found in cool-temperate, mixed-deciduous broad-leaved forests, in mixed-deciduous broad-leaved and coniferous forests, and in nearly pure coniferous forests at high altitudes or latitudes. On Mount Jinfu Shan in southeastern Sichuan Province, D. sinensis is sometimes found growing in bamboo thickets. Plants characteristically associated with Diphylleia in each of the three disjunct regions are listed in TABLE |. Hara (1959), Numata ef al. (1972), and Numata (1974), without referring specifically to D. grayi, have discussed some of the associa- tions in which it occurs in Jap e to 1ts use as a ocak plant and to the destruction of its habitat, Diphylleia sinensis is becoming increasingly scarce. In Japan and the United States, however, large areas where Diphylleia grows are under protection in national forests and national parks, and the plants are in no danger of extir- pation. ECONOMIC IMPORTANCE In China, where Diphylleia sinensis has long been used as a folk remedy, the rhizomes are collected during the summer or fall, cut into slices, and dried in the sun. An infusion of the rhizomes has been employed in the treatment of injuries from falls; chewing and swallowing the rhizome has been a remedy for stomach trouble. The rhizome, however, contains poisonous substances, and care should be taken in using it; it should not be taken during pregnancy (Fu. 76; Anonymous, 1974). It has also been reported (Anonymous, 1974) that the rhizome has been employed in treating lumbago, rheumarthritis, ““men- oxenia,” and gynecologic ailments, and that the roots are useful in lowering fever, detoxification, dissolving blood clots, and improving circulation. Diphylleia grayi is occasionally cultivated as an ornamental plant in Japan. Lloyd and Lloyd (1887, p. 120), commenting on D. cymosa, stated “It is not an article of commerce, there being no demand for it.” CHEMISTRY Murakami and Matsushima (1961) investigated chemical compounds in the rhizomes of Diphyileia grayi. They reported podophyllotoxin, picropodophyl- 1984] YING ET AL., DIPHYLLEIA ay lin, 8-apopicropodophyllin, kaempferol, quercetin, diphyllin, and an additional substance that they assumed to have a 4-arylnaphthalene skeleton. They be- lieved that the constituents picropodophyllin and 8-apopicropodophyllin were formed from podophyllotoxin during separation. Lloyd and Lloyd (1887) noted that after evaporation, alcohol extractions of the rhizomes of Diphylleia cymosa produced a bitterish, acrid-tasting resin similar to that obtained from Podophyllum peltatum. They also reported that Diphylleia extracts had no medicinal properties, in contrast to those of Podo- phyllum, and (p. 120) “careful investigations demonstrated that no trace of an alkaloid or other interesting constituent existed in the plant.” More recent investigations by Kimura (1963) revealed that extracts of Di- phylleia grayi contain substances similar to colchicine and podophyllin that produce an antimitotic effect on cancer cells of MTK-sarcoma III, Yoshida- sarcoma, and Ehrlich ascites carcinoma: the cells die without progressing be- yond metaphase, thereby causing regression in tumors. Toyokuni and Toyokun1 (1964) used this evidence as a basis for a classification of the Podophyllaceae in which they recognized two subfamilies. One, the Podophylloideae, contains podophyllin or podophyllinlike substances that inhibit tumors; the other, the Glaucidioideae, contains no podophyllinlike substances and has no effect on tumors. They considered the Podophylloideae to contain two genera, Podo- phyllum (nine species) and Diphylleia (D. cymosa and D. grayi). The sole member of their Glaucidioideae 1s Glaucidium palmatum Sieb. & Zucc. ANATOMY Material used in this study consisted of flowers, buds, and vegetative parts of Diphylleia cvmosa and D. grayi, and fruiting peduncles and pedicels of D. sinensis, all preserved in FAA. Voucher specimens are cited in APPENDIX 1. Methodology was the same as in Terabayashi (1983). For a more detailed description of the anatomy of Diphy/leia, that paper should be consulted. RHIZOME AND Root Kumazawa (1930) discussed the anatomy of the rhizome and root in Di- phylleia grayi. Our study confirms Kumazawa’s findings and shows the root and rhizome anatomy of D. cymosa and D. grayi to be similar. Secondary xylem elements and an interfascicular cambium are not produced, and the phellogen in the rhizome is subepidermal in origin. The roots of Diphylleia are adventitious and about | mm in diameter. The stele of the roots in D. grayi and D. cymosa is usually tetrarch (see FiGure |), with secondary xylem ele- ments rarely produced. According to Zhong Guo Yixui Gexul Yuan Yowu Lianjuoso (1979) the root of D. sinensis differs from those of the other two species in having a pentarch stele. STEM Studies of Harvey-Gibson and Horsman (1919), Himmelbaur (1913), Ku- mazawa (1930), and Worsdell (1908) showed that Diphy//eia is among a small number of dicotyledonous genera having the monocotyledonous feature of 60 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Taste 1. Plants characteristically associated with Diphylleia.* CHINA JAPAN UNITED STATES D. sinensis D. grayit D. cymosa Abelia dielsii Abies fargesit Abies mairesil Abies fraseri Abies veitchii Acer caudatum Acer rufinerve Acer pensylvanicum var. multiserratum Acer tschonoskii Acer saccharum Acer tetramerum Acer yee NSE Acer spicatum Actaea asiatica Actaea astatica Actaea pachypoda Aesculus octandra Arisaema triphyllum Asarum caudigerum Athyrium fallaciosum Athyrium spp. Athyrium asplenioides {rhyrium filix-femina Betula albo-sinensis Betula ermanti Betula aa var. septentrionalis Betula les Betula fargesii Cacalia roborowskii Cacalia adenostvloides Cacalia muhle ie Cardamine engleriana Cardamine clematitis Caulophyllum robustum Caulophylhium robustum Caulophyllum aes Chrysosplenium micro- spermum Cimicifuga foetida Cimicifuga racemosa Circaea ao DHA Circaea alpina Circaea alpina subsp. imaicola subsp. alpina subsp. al/pin Clintonia ene Clintonia udensis Clintonia umbellilata Coptis trifolia Disporum lanuginosum Dryopteris sinofibrillosa Dryopteris austriaca Dryopteris austriaca es crash zoma Dryopteris intermedia “agus grandifolia eae piles ae Kingdonia uniflora Kinugasa japonica Ligularia stenocephala Lindera umbellata Lindera benzoin Liriodendron ee Liriodendron tulipifera Lonicera kungea Magnolia acuminata Magnolia fraseri Oxalis acetosella Oxalis acetosella Panax pseudo-ginseng var. bipinnatifidus Paris thibetica Paris polyphylla Paris tetraphylla Peracarpa carnosa Peracarpa carnosa . circaeoides var. a Picea wilsonii Picea Jezoensis Picea rubens een matsumureana Polygonatum roseum Polygonatum pubescen Polystic hum retrorso- Polystichum ee x paleaceu} Prunus gracilifolia Prunus grayvana Prunus serotina Pseudocystopteris spinulosa 1984] YING ET AL., DIPHYLLEIA 61 TABLE 1 (continued). CHINA JAPAN UNITED STATES D. sinensis D. grayit D. cymosa Rhododendron concinnum Rhododendron maximum Rhododendron fargesti Rhododendron catawbiense Rhododendron purdomii Ribes glaciale Ribes rotundifolia Rosa omeiensis Salix cathayana Saxifraga micranthidifolia Schisandra incarnata Smilacina henryi Smilacina racemosa Smilacina japonica Smilacina japonica Sorbus koehneana Streptopus Ca Streptopus roseus var. japon Thalictrum ichangense Thalictrum clavatum Thalictrum przewalskii Tiarella polyphylla Tiarella polyphylla Tiarella cordifolia subsp. cordifolia Tilia americana (incl. 7. heterophylla) Trautvetteria japonica Trautvetter ig caroliniana Trillium tschonoskii Trillium smallit Trillium erectum Trillium See Trillium vaseyi BG Soest aaa chin Tsuga diversifolia Tsuga canadensis alo aan Vaccinium japonicum Vaccinium erythrocarpum r. sinensis var. japonicum V ‘iburni num betulifolium Viola acuminata * Author names have been omitted to save + Partially from Hara (1959), Numata aoe ne Numata er al. (1972). scattered vascular bundles in the stem stele. The aerial stem in all three species is about 10 mm in diameter and contains 70 to 80 scattered vascular bundles in the portion below the leaves: between 40 and 50 vascular bundles occur in the internodal region (see FiGure 4). The interior bundles are larger and mature earlier than those near the periphery. A sclerenchymatous ring, in which the phloem of the outer small bundles 1s embedded, is found near the periphery of the mature aerial stem. Kumazawa (1930, 1932) reported that the inner bundles give rise to the leaf traces in D. grayvi. The same is true in D. cymosa. After the separation of traces to the upper leaf, a rearrangement of the vascular bundles occurs so that there are two concentric circles of vascular bundles in the peduncle. This configuration persists through the pedicel in D. grayi and D. sinensis, but in D. cymosa one circle is lost during branching of the inflo- rescence, so the stele of the pedicel consists of only a single circle of vascular bundles (FIGURE 3). Where two circles of vascular bundles occur, the bundles 62 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Figures 1-3. Cross ee cae 1, Diphylleta gray, root, showing tetrarch stele (EN. hl ith; XY, Perens PE, pericycle; , phloem; PI, xylem; scale = 0.1 mm). 2, D. gray, vascular bundle in stem ee metaxylem (sl Aly immature in this sample): PX. protoxylem; PH, protophloem; CA, inactive cambium: scale = 0.1 mm). 3, vasculature f peduncles (below) and pedicels (above), scale = 1 mm: a, D. eer single circle of vascular bundles in pedicel): b, D. gravi c. D. sinensis. in the inner circle are larger than those of the outer. A cross section of a single vascular bundle in the stem of D. gray is shown in FiGure 2. LEAF The petiole of Diphylleia is stemlike, with scattered vascular bundles. There are 40 to 50 vascular bundles in the petiole of the lower leaf, and 20 to 30 in 1984] YING ET AL., DIPHYLLEIA 63 Ficure 4. Cross sections of aerial shoots of Diphylleia showing vasculature: a-d, D. cymosa, e-h, D. grayi (note sessile upper leaf in h). a and e, near ground level; b and f, just above node of lower leaf: ¢ and g, internodal region; d and h, just above upper leaf. Scale = that of the upper leaf. The petioles are round in cross section from about the middle upward, but the lower portion shows dorsiventrality in both external and internal morphology (FIGUR The anatomy of the leaf blades in Diphylleia grayi and D. cymosa was studied by Oganozova (1974), who found the mesophyll of D. cymosa to be ca. 0.07 mm thick, with the palisade layer one cell thick and the spongy tissue four 64 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 cells thick. He reported the number of stomata of 7 lower epidermis to be 25 per mm? in D. cymosa and 24 per mm? in D. gra The leaves are folded umbrellalike in bud in Diphy ioe as they are in Podo- phyllum and Dysosma Woodson. Kumazawa (1937) indicated that in Podo- phyllum and Dysosma the summit of the folded leaf is at the position of attachment to the petiole, although in Diphylieia the fold in the leaf is above the point of attachment. FLOWER Terabayashi (1983) examined the floral vasculature of Diphylleia cymosa and D. grayi in detail. Unfortunately, in both his study and ours flowering material of D. sinensis was unavailable for comparison. In D. grayi the double circle of vascular bundles in the pedicel continues into the receptacle; the inner circle contains the main bundles. In D. cymosa, where the pedicellate stele is composed ofa single circle of vascular bundles, an outer circle of bundles arises from the stelar bundles at the lowest level of the receptacle, so the receptacle contains the two circles of bundles as in the peduncle and stem. No major differences are seen in the trace patterns to the sepals, petals, stamens, and pistils in the two species. The traces to the outer sepals originate mainly from the outer circle of vascular bundles, while those to the inner sepals, petals, and stamens usually derive from adjacent bundles in the inner circle. In exceptional cases, weak bundles from the outer circle may fuse with the petal traces. The pistil is traversed by a few ventral bundles and a dorsal bundle that extends to the stigmatic region. The ovary wall 1s covered by a network of veins. The ovular traces arise from ventral bundles, while ie in the placental region originate from bundles other than ventral bundle Chapman (1936) and Kaute (1963) studied oistil anatomy in Diphylleia and interpreted the pistil as being pseudomonomerous. Kaute (1963) observed abnormally dimerous pistils with a single locule but with two placentas in D. cymosa. Such abnormally dimerous pistils were also observed in D. grayi by Terabayashi in this study. The monomerous pistil in Diphyl/eia may have been derived from a dimerous pistil through reduction. EMBRYO In Diphylleia cvmosa the ovule is bitegmic and crassinucellate: embryo sac formation is of the Polygonum type, and pollen formation is simultaneous (Mauritzon, 1936). Terabayashi found the ovular morphology and embryo sac formation in D. grayi to be the same as in D. cymosa. The seed coat of D. grayi is exotestal, according to Corner’s terminology (Corner, 1976; Takhtajan & Melikian, 1972). SEEDLING The seedlings of Diphylleia are characterized by long “‘cotyledonary tubes” (Himmelbaur, 1913; Terabayashi, in prep.). The cotyledon is two-parted, with the “cotyledonary tube” and an orbicular lamina. The plumule is hypogeal. The seedling does not produce foliage leaves during the first year after ger- mination. The stele in the radicle and hypocotyl is diarch. 1984] YING ET AL., DIPHYLLEIA 65 CYTOLOGY The number and karyotype of the chromosomes of Diphyilleia gravi are well known. Counts of 2” = 12 have been reported in this species by Kurita (1956), Kuroki (1967), Matsuura and Suto (1935), Miyagi (1930), S. Noda and Fuji- mura (1970), Noguchi and Kawano (1974), and Soeda (1942). The anomalous count of 2 = 16 attributed to Lee (1967) is in error. Lee reported 2” = 16 for Caulophyllum robustum Maximowicz; this count was later mistakenly reported as being for D. grayi by Noguchi and Kawano (1974). As far as we have been able to determine, Lee did not examine chromosomes in Diphylleia. The single report (Langlet, 1928) of 2n = 12 for Diphylleia cymosa was determined from plants in cultivation in Europe. Langlet considered Diphylleia to be monotypic, with D. cymosa as the sole species. Although it can be assumed that the material he was working with—and probably the species in cultivation in Europe at the time—was D. cymosa, there is no way to prove this. Langlet did not cite vouchers for his cytological studies, and we saw no specimens collected or annotated by him among those that we examined. Until Langlet’s report can be verified, it seems best not to attribute counts to D. cymosa. No counts have been made of the Chinese Diphylleia sinensis. Chromosome counts of Diphylleia are summarized in TABLE 2. Kurita (1956), Kuroki (1967), S. Noda and Fujimura (1970), and Noguchi and Kawano (1974) reported similar karyotypes in plants of D. grayi collected from widely separated populations in Hyogo, Iwate, and Tochigi prefectures, Japan. All reported two pairs of large (length 10.7-15.5 um; all measurements from Kurita, 1956, and Kuroki, 1967), metacentric chromosomes, one pair of medium (length 9-11.1 um), submetacentric chromosomes, and one pair of medium (length 8.4-10.8 wm), submetacentric chromosomes with a constric- tion at the midpoint of the shorter arm (Noguchi and Kawano (1974) reported the constriction to be at the distal end of the short arm, but their illustration shows it to be more nearly medial). Of the two remaining pairs, all of the authors reported them as being small (length 6.3-7.6 um), with one pair being telocentric and the other having a subterminal constriction. Soeda (1942) found an additional constriction on the short arm of the small, telocentric chromo- some, but none of the other authors mentioned this. S. Noda and Fujimura (1970) also examined mitosis in developing seeds. In the plants in their study, they found that fruit-set was about 60 percent, but only 21.1 percent of the ovules developed into mature seeds. They (pp. 2549, 2551) attributed this high sterility to an“... asynchronization of mitosis” that they observed in endosperm tissue. PALYNOLOGY'! Pollen samples were collected from unopened buds or young flowers on herbarium specimens (see APPENDIX 2 for vouchers) of three sheets each of Diphylleia cvmosa, D. grayi, and D. sinensis. The samples were acetolyzed ‘Contributed by Masamichi Takahashi, Biological Institute, Faculty of Education, Kagawa Uni- versity, Takamatsu-shi 760, Japan. Tas_e 2. Reported chromosome counts for Diphylleia. CHROMOSOME TAXON LOCALITY REFERENCE VOUCHER* D. cymosa 2n= 12 U.S. A.t Langlet, 1928 S D. grayi 2n= 12 see Hes forest in mountains Miyagi, 1930 KIEL fN n=6 Japan Hokkaido, Ishikari Shicho, Matsuura & Suto, 1935 SAP . Mol 2n= 12 aoe Ho kkaido Soeda, 1942 SAPS 2n= 12 Japan, Honshu, Tochigi Pref., Kurita, 1956 Ehime Univ. Mt. Shirane 2n= 12 Japan. Honshu, Iwate Pref.. Kuroki, 1967 Ehime Univ. Hachimantai 2n= 12 Japan, Honshu. Hyogo Pref.. S. Noda & Fujimura, 1970 ? Hyono-sen 2n= 12 Japan, Honshu, Tochigi Pref., Noguchi & Kawano, 1974 Toyama Univ. Nikko * No voucher specimens were cited in any of the reports. If vouchers exist. they are probably in the herbaria indicated. + See text. WOLAYOUUY GTIONYV FHL JO TWNUNOL 69 “10A] 1984] YING ET AL., DIPHYLLEIA 67 Figures 5-8. SEMs of pollen of Diphyilleia, all x 1400: 5, D. cymosa (J. K. Small 5.n., 28-29 May 1892), polar view; 6, 8, D. sinensis (Licent 5325), polar and equatorial views; 7, D. grayi (3 July 1952, A. Kimura & S. Sugaya s.n., 3 July 1952), inaperturate. according to procedures outlined in Erdtman (1960), dehydrated in ethanol, transferred to amyl acetate, and critical-point dried. They were then coated with gold by using an ion sputtering apparatus (Damblon, 1975) and examined with an Akashi MSM-101 scanning electron microscope. For transmission electron microscopy (TEM) the pollen was fixed in | percent osmium tetroxide, dehydrated in an ethanol-propylene oxide series, and embedded in a low-viscosity Epon mixture according to the method of Spurr (1969). Glass knives were used for sectioning on an LKB ultratome. The un- stained — were examined with a Hitachi H-500 transmission electron microsco Pollen of Diphylleia has been examined by Kumazawa (1936) and Ikuse (1956), who used light microscopy, and by Nowicke and Skvarla (1981), who used light microscopy and scanning and transmission electron microscopy, but none of these authors examined all three species. Diphylleia 1s unique in the Berberidaceae s./. in having spinose pollen grains. Pollen features support other —’” 68 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Figures 9-12. Pollen aie ornamentation in Diphylleia. 9, TEM, D. tes (77. Takahashi 366), section through spine, x 12,000. 10-12, SEMs, all x 3670: 10, D. cy- he (Kologiski & Perino 165), r .D. grayi (A. Kimura & S. Sugaya s.n., 3 July os , D. sinensis (Nan-shui-Bet- Aigo Team 7 16+¢ data that indicate a closer relationship between D. cymosa and D. sinensis than between either of these two and D. grayi. In gross morphology, pollen of D. cymosa and that of D. sinensis are basically the same (Ficures 5, 6, 8): both are tricolpate or rarely hexarugulate. In contrast, the pollen of D. gravi (FIGURE 7) has irregular apertures, which is the same as that reported by Nowicke and Skvarla (1981) in several species of Berberis L. and Mahonia Nutt. Kumazawa (1936) reported finding some pollen grains of D. grayi that had less densely staining spineless, “round areas” containing densely staining “punctations” that he regarded as the germinating pores, while Ikuse (1956) found pollen of D. grayi to be polyforate, 34-36 um in diameter, and with spines 2.5 um long. In exine ornamentation, however, D. cyimosa is more similar to D. grayi than to D. sinensis. The exine of the first two species is characterized by stout spines 1.5—4.5 wm long, while that of D. sinensis has slender spines 0.5-2.8 um long. This difference between D. cymosa and D. sinensis was also noted by Nowicke and Skvarla (1981). FiGures 9-12 show exine ornamentation of the pollen of the three species, while TABLe 3 gives pollen size of the samples examined in this study. TAXONOMIC HISTORY Diphylleia was first discovered by André Michaux in North Carolina and was described and illustrated in his F/ora Boreali-Americana (1803). Plants of 1984] YING ET AL., DIPHYLLEIA 69 Tasce 3. Pollen size in Diphylleia.* SPINI Pol LEN DIAMETER LENGTH COLLECTION um) (um) D. cymosa Kologiski & oe 165 36.7 + 2.5 x 32.6 + 2.7 1.5-4.5 J. K. Small s.n., 32.5 + 2.5 x 29.7 + 2.0 1.5-3.7 28-29 May “392 Radford 5239 35.8 + 2.4 x 31.1 + 2.3 1.5-3.8 D. grayi Takahashi 366 34.9 + 2.7 x 31.5 + 2.2 1.5-4.2 Kimura & Sugaya S.n., 37.7 + 1.7 x 33.3 + 2.2 1.5-4.7 3 July 1952 Saito & Kaneko s.n., 35.0 + 2.2 x 30.5 + 2.5 1.4-4 18 May 1961 D. sinensis Delavay 3862 31.3 + 2.3 x 30.2 + 1.7 0.5—2.0 Nan-shui-Bei-diao Team 7164 30.1 + 2.5 x 28.5 + 2.6 0.7-2.5 Licent 5 29.6 + 2.1 x 28.4 + 2.0 0.5-2.8 * Thirty pollen grains were measured from each specimen. Diphylleia from Japan and Sakhalin were probably first collected around 1855 by C. Wright and J. Small; Asa Gray (1859) considered them to be conspecific with the North American D. cymosa—an opinion shared by Diels (1900). Schmidt (1868), however, recognizing the plants from Sakhalin to be different from Michaux’s D. cymosa, described them as D. grayi, and Kitamura and Murata (1962) subsequently treated them as a subspecies of D. cymosa. Au- gustine Henry apparently first collected D. sinensis in 1888 in what is now the Shennongjia Forest District in northwestern Hubei Province (Henry im Thi- selton-Dyer, 1889), but he believed the plants he found to be D. gray. Authors subsequent to Henry treated these plants from central China as either D. cymosa or D. grayi until 1947, when Li described them as D. sinensis. Since the characters that can be used to separate the three taxa are discon- tinuous, we prefer to treat them as three distinct species. RELATIONSHIPS AND PLACEMENT OF DIPHYLLEIA The familial placement of Diphylleia has been controversial. Schultz (1832) was the first to place Diphylleia in a separate family, the Diphylleiaceae (in which he also placed Sarracenia!), while Tischler (1902) proposed that the two genera Diphylleia and Podophyllum (which included the then-undescribed Dys- osma) represented the family Podophyllaceae. In this, Tischler has been fol- lowed by Hutchinson (1969), Takhtajan (1969), and Airy Shaw (1965), although Hutchinson and Airy Shaw expanded the concept of the Podophyllaceae to include most or all of the herbaceous genera of Berberidaceae. Engler (1903) established a new subfamily in the Berberidaceae, the Podophylloideae, in which he placed Diphyilleia and all other herbaceous members of the family. 70 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 His treatment was followed by Ernst (1964) and Thorne (1968). Cronquist (1968, p. 365) placed the Podophyllaceae (presumably including Podophyllum and Diphylleia) in his Ranunculaceae but later (1981, p. 126) returned them to the Berberidaceae. Despite the controversy that exists over the familial placement of Diphylleia, there has been little doubt concerning its closest generic relatives. Based on chromosomal, anatomical, and chemical evidence, as well as on overall mor- phological similarity, Diphylleia unquestionably has its greatest affinities with Podophyllum and Dysosma. Meacham’s (1980) study supports this view. How- ever, Nowicke and Skvarla (1981), using palynological evidence, do not feel that these genera are particularly close. POSSIBILITIES FOR FURTHER STUDIES Perhaps one of the most interesting studies that could be undertaken with Diphylleia would be one dealing with reproductive biology. As far as we have been able to determine, there have been no investigations of this topic. The spinose pollen grains (like those of the Compositae) and the showy clusters of flowers that are held high above the forest floor in early spring strongly suggest insect pollination. However, as in many other herbaceous plants of mixed deciduous forests, the extensive rhizome system also suggests a degree of veg- etative reproduction that might account for the large colonies encountered in D. cymosa and D. grayi. The plants occurring singly or in open colonies in D. sinensis possibly indicate a greater dependence on reproduction by seed than by vegetative means, especially since individuals are often found several meters apart. In connection with reproductive biology, it would be of interest to know whether plants are self compatible and, if so, to what extent outcrossing takes place. Study of inter- and intraspecific crosses and knowledge of chromosome behavior in any resultant hybrids are also desirable. Chromosome counts and karyotype analyses of D. cymosa and D. sinensis would enable comparison with the cytologically well-known D. gravi. Chemical analysis and comparison of the three species could also provide useful information. Except for the above-mentioned work of Murakami and Matsushima (1961), Kimura (1963), and Toyokuni and Toyokuni (1964), we are unaware of any chemical studies involving Diphyvileia. Comparative fla- vonoid studies might prove useful, especially in light of the differences among the three taxa revealed by other comparative studies. Comparative evidence from any of the above areas would improve our understanding of the rela- tionships of the three species and could also make possible more detailed comparisons between Diphy/leia and other genera of the Berberidaceae. TAXONOMIC TREATMENT Diphylleia Michaux, Fl. Bor.-Amer. 1: 203. pls. 19, 20. 1803. Perennial herbs with thickened, creeping rhizomes and coarse, fibrous roots, the rhizomes formed of distinct annual increments, producing stout, 2- (rarely 1984] YING ET AL., DIPHYLLEIA 7\ 3-)leaved stem that separates at base in autumn along marked articulation, leaving broad, bowllike excavation on rhizome. Leaves alternate; blade peti- olate and peltate (except upper leaf in Diphylleia grayi, which 1s sessile or subsessile and attached at sinus), transversely oblong to reniform-orbicular, 2-cleft with divisions shallowly to coarsely lobed and prominently dentate, palmately veined with main veins connected by secondary, reticulate veins, pubescent or sparsely pubescent with unicellular hairs. Inflorescence a terminal, pedunculate (peduncle rarely branched below), usually many-flowered cyme or umbel, the branches glabrous or pubescent. Flowers pedicellate, actinomorphic, 3-merous; sepals 6 in 2 (rarely 3) whorls, white or pale green, deciduous prior to or just at anthesis; petals 6 in 2 whorls, white; stamens 6, antipetalous, the anthers basifixed, longitudinally dehiscent, the thecae separating from connec- tive and ultimately attached only at apex, the pollen conspicuously spiny; ovary superior, ellipsoid, unilocular, the style absent or very short and thickened, the stigma peltate, cristate, the placentation parietal near base of ovary, the ovules to 11, anatropous. Fruit a globular or broadly ellipsoid berry, dark blue, glaucous, to | cm broad, borne on thickened, disclike, lobed receptacle. Seeds oblong to ovoid, rounded on all sides or flattened to concave ventrally, straight or slightly curved, surface minutely rippled and microscopically striate. Embryo straight or slightly curved, ca. 2-4 mm long, with 2 cotyledons, embedded in endosperm. Chromosome number: n = 6. Type species. Diphylleia cymosa Michaux. A genus of three species, with one in each of three widely disjunct areas: the southern Appalachian Mountains of the southeastern United States; central and northern Japan, Sakhalin, and the southern Kuril Islands (Kunashiri Is- land); and central and southwestern China. Key TO THE SPECIES OF DIPHYLLEIA 1. Inflorescence glabrous; ovules 2 to 4; leaves glabrous or sparsely eget ee sesig ih arehytds Bese Set Sa 4 as ce sate ea gsra cute tte. iatepee mata atne Mea dives Ace esa ir ane Lehane . D. cymosa. 1. Inflorescence pubescent; ovules 5 to 11; leaves pubesce 2. Upper leaf distinctly petiolate, the petiole more than . 5 cm long; ae s less than Dz: 9 mm long and 6 mm wide; flowers usually 15 or more. ........ 2. sinensis. 2. Upper leaf sessile or with petiole less than 1 cm -_ petals more than 8 m long and 6 mm wide; flowers usually 12 or fewer. ............--5 3. D. grayi. 1. Diphylleia cymosa Michaux, Fl. Bor.-Amer. 1: 203. 1803. FiGure 13. Plants 4-12 dm tall, growing in dense or loose colonies, occasionally as scattered individuals. Petiole 7-22 cm long in lower leaf, 3-15 cm long in upper; blade peltate, suborbicular, reniform or transversely oblong, deeply cleft at apex and base, 14-30 by 19-47 cm in lower leaf and 9.5-33 by 13-42 cm in upper, deeply cut to lobed at margin and coarsely doubly serrate, sparsely pubescent along main veins beneath or glabrescent, usually glabrous above or with few hairs along main veins. Peduncle solitary, occasionally bilurcate, 0.6— 3.3 cm long, glabrous; inflorescence 7.5—41 cm long including peduncle, 3.7- dz JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 ig? \\ fia | Ro YZ ia } Figure 13. Diphylleia cymosa (based on Chinese-Amer. Exped. 853, pv): a, upper leaf, inflorescence, rhizome, roots; b, flower; c, outer sepal: d, inner sepal: e, outer petal: f, inner petal; g, stamen: h, ovary; 1, fruit. 1984] YING ET AL., DIPHYLLEIA 73 14 cm wide, glabrous (rarely with few scattered hairs when young). Flowers (6 to) 14 to 70 or more; pedicels 0.8—4 cm long; outer sepals lanceolate, oblan- ceolate to spatulate, or reduced and scalelike, ca. 1.7-4.5 by 0.4 mm; inner sepals obovate to suborbicular, 2.5-6 by 3.5-4 mm; outer petals narrowly obovate, 9-11 by 4.5-6 mm; inner petals elliptic-obovate to obovate, 10-13 by 6-7 mm; stamens 3.6—4.5 mm long, filaments 1-2 mm long, anthers ca. 2 by 1.2 mm; ovary ellipsoid, 3.5—5 by 2—2.5 mm, style ca. 0.3 mm long, stigma ca. 0.5 by | mm. Berry globular to very broadly ellipsoid, 0.5-1.2 cm broad, blue, glaucous, on pedicel 1-3.8 cm long. Seeds 2 to 4, + oblong, 3.9-7 by 2- 4.7 mm, rounded dorsally, flattened to concave ventrally. Chromosome num- ber: n = 6 (but see under cytology above). Tyre. United States, mountains of North Carolina, 1786?, A. Michaux s.n. (lectotype, Pp, photo A; isolectotype, P). DISTRIBUTION AND ECOLOGY. Moist places in mixed deciduous and coniferous forests, often along seey and small streams, |000-1650 malt. United States: southwestern Virginia, western North Carolina, and eastern Tennessee to north- eastern Georgia and extreme northwestern Oconee County, South Carolina; restricted to Blue Ridge of southern Appalachian Mountains. (Map 1.) SPECIMENS EXAMINED. United States. GEorGIA. Rabun Co.: Rabun Bald, 34°57'N, 83°18'W, Boufford & E. W. Oe 17033 (NcuU); along Darnell’s Creek SE of Dillard, Duncan & McDowell 10852 (Ga); 2 mi N of Clayton, Pyron & aes 847 (GA). Towns Co.: N-facing slope, Hogpen Gap, Richard Russell Scenic Hwy., Colle 346 (NCU z N- costae ravine just N of summit of Hightower Bald, Duncan 7597 (c A). Union Co.: i W of Vogel State Park, Hardin 259 (Ga); Brasstown Bald, Howell 407 (us); Sosbee aoe ae Vogel State Park, Kral 59676 (vps); |.5 mi from top of Brasstown Bald, Lord 364 (NcU); cove E of Wolfpen Gap, Pyron & Duncan 46 (GA); Blairsville, Lock Gap, /. Stuckey s.n. (vpB). White Co.: 2.5 map mi above Jct. of Chattahoochee R. and Spoibine Creek, Duncan & Hardin 16361 (Ga); Hogpen ie offhwy. 348, Wohlers 122 (GA). NORTH CAROLINA. “Mountains of North Carolina,” 4A. Michaux s.n., 1786? (Michaux herb. at p, IDC 6211. 47: I. 2, 3). Iron Mt., Canby s.n., pee 1876 (ny). Ashe Co.: Bluff Mt., Radford 44012 (Ncu). Avery Co.: 1.2 mi pee of Minneapolis on U.S. 19E, Ahles & Ashworth 3945] (Ncu); NNW of Linville, 1 mi SSW of jet. N.C. 184 and 105 on 105, Ahles & Duke 43632 (NCU); ns Mt., Blomquist 3681 KE), Heller s.n., 22 July 1890 (LE, NY), Mohr s.n., 28 July 1894 . s), Mullen 81 (NCSC) heal es S& 13 (DUKE); ca. 7 mi ENE of Linville along U.S. route 221, McVaugh 8937 (GA, NCU, vpB); 2 mi E of Linville along U.S. route 221, ae & Umber s.n., 3 June 1973 on vic. of Banner Steele 18 ie s); Linville Falls, Weiss s.n., 30 May 1959 (ver); NE of Linville, 3 km E of U.S. route 221 on county road 1514 (Edgemont Road), £. HW. Wood & se ies 113 — (BM, CM, KYO). Buncombe Co.: Craggy Mt.. Biltmore Herb, 1213° (BM, GH, L, Us, Ww); Pisgah a Forest, ca. [O mi SE of Dillingham, Boufford et al. ee ee U); s flank Black Mt., NV. L. & Mrs. Britton s.n., 18 Sept. 1885 (Nv); Black Mt., Canby s.n., June 1868 (Ny): sie Ridge Pkwy. at Craggy Gardens. Freeman 58196 Nee wet woods in Craggy Gardens, Hicks & Bartley nave a Black Mts., LeRoy s.n. (Nv), Rugel 243 (pM); Graybeard Mt., Pollard s.n., 16 May 1901 (us); Cries Mts., Rvdberg 9446 (LE, NY). Burke Co.: near Linville, Barkley & hie 2031 (NY, US). Caldwe ll Co.: NW corner of Caldwell Co. on U.S. 221, hwy. to Grandfather Mt., le & Stewart 1575 (Ncw). Cherokee Co.: 2.3 mi E of Tennessee state line on Tellico R. Road, Pittillo & Floyd 5710 (wcuH). Clay Co.: near Perry Gap, ravine at end of U.S. Forest Service Road along Buck Creek (N of U.S. route 64), Boufford ¢ el a 14165 (wcu), 1700] (Bm); ca. 15 mi E of 74 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Haysville, Chunky Gal by U.S. route 64, Kral 60290 (vps); Chunky Gal Mt., W of Rainbow Springs on U.S. 64, Pittillo et al. 4289 (wcun), Buck Creek near U. S. 64, 2 mi SW of Clay-Macon Co. line, Radford 12162 (Ncu), Perry Gap Road above Buck Creek, 4miN of US. 64, Fox & Godfrey cs (Ncsc). Graham Co.: 5.1 mi N of jet. of Cherokee Co. roads 1391 and 1399 on U'S. Forest Service Road 423, Boufford et al. 14518 (Ncu): 4 mi E of Fontana, Radford 11870 ee u, vps); 4 mi S of Sonne Radford 14196 (ncu); Joyce Kilmer Mem. Forest, Tucker & Pittillo 141 (wcun). Hayw ood Co.: Junaluska Mt., Batchelder S081, 5082 (Gu): ee uska Mt., Lake Junaluska, Blomauist 3682 (DUKE); ca. 1.6 mi S of Sunburst on N.C. 215 along West Fork Pigeon R., Boufford et al. 15041 (ncu); Mt. Sterling, Caughey a (ncsc); Bill Camp Cove, Couch s.n., 24 July 1926 (Ncu); Jonathan Creek, Pyron 63 (GA): Eagles Nest, Ruth 367 (Gu). Blue Ridge Pkwy. at Frying Pan Gap, Smathers s.n., 25 June 1958, 22 June 1959 (wcuH); Balsam Mt., J. D. Smith s.n., 11 Aug. 1882 (us); near Waynesville, Eagles Nest, Standley 5459 (us); county road 1334 2.8 mi N of intersection with county road 1335, Wyatt 267 (puke); Crabtree Bald, county road 1505 ca. 5 mi NE of intersection with N.C. route 209, H’yart et ii 598 (puUKE). Jackson Co.: 14.7 mi NW of N.C. route 215 on Blue "Ridge Pkwy., Boufford et al. 13529 (Ncu); Balsam, Braun s.n., 14 July 1911 (us), 5.2 mi S of Dillsboro on county road 1371, Bryson 47 (wcun); Tuckaseegee R., 12 way from Sylva to Cashiers, Coker s.n., 21 Aug. 1939 (Ncu); 10 mi S of Cullowhee P.O. on N.C. 107, Crisp s.n., 29 June 1965 (wcun); E of Glenville, Slatten Creek, Godfrey & O'Connell my (DUKE, nesc): Soco Falls, Hendrix B-22 (puKe); 14 mi SSE of Cullowhee P.O. on N.C. route 107, Hoffman 70 (wcun); Mull Creek, Murtagh 230 (wcu a Nantahala Natl Forest, N.C. route 107 at pipeline from Lake Thorpe, Pittillo 2876 (©, FARM, GA, GH, NCU, NY, vpB, WCUH); Rough Butt Bald Mt., Ramseur 389 (NCU); Wet Camp Gap, W”. B. & M. B. Schofield 8527 (puke); ca. 4 mi S of Thorpe Power Sta. on N.C. route 107, Sharpe 26 (wcun); above Glenville Power Plant, Smith $.n., ae May 1953 (weun); 15.1 m1 S of Cullowhee P.O. on N.C. route 107, Taylor 38 (wcun). Macon Co.: Satulah, L. B s.n., 24 May 1934 (Ncw); near a. fore Herb 1213 (GH, LE, NY, P, US, W), 1213° (FI, G); under Dry Falls, Coker & party s.n., 28 Aug. 1932 (Nc a near Rainbow Springs, D. S. Correll 3522 (puke); 4.5 mi W of Fish Checking Sta. on Nantahala Forest Road, Fox & Godfrey 3081 (csc); U.S. route 64 at Glade Gap, Freeman 59143 (Ncw), Highlands, Gibbes s.n., July 1882 (xy), Harbison s.n., 13 July 1901 (Gu), county road she 2.2 mi from jet. with county road 1679, Metheney 076 (wcun); Cullasaja R. at y Falls, Oosting 34427 (puke), Highlands, Palmer 42502 (Ny), county road 1001 S$ ree tae Gap to Wildcat Road, E to Bryson Branch, Pittillo 4050 (wcun), 13 mi N of U.S. route 64 on Rainbow Springs-Aquone Forestry Road, Radford 5239 (Ncu), 4 mi N of Highlands, Cole Mt. Road, Radford 6123 (Ncu); Cullasaja Gorge, High Falls, W. B. Schofield 8776 (puke); Crow Creek ca. 0.5 mi below Cullasaja Falls, Stewart & Hechenbleikner s.n., 25 July 1938 (Ncu); Stewart Trail aie Bearpen and = bert Mt., Stewart & ciate ie sm, 12 Aug. 1938 (Ncu). Madison Co.: 0.2 mi N of Betsy Gap and Haywood Co. line on N.C. 209, Bozeman et al. 316 J2 (GA, GH, NCU, NY, VDB). McDowell Co.: near Big Cr raggy on Blue Reet Pkwy., Ahles & Bell 17697 (Ncu); Blue Ridge, W. F. s.n., 1872 (GH). Mitchell Co.: an Mt., Cin. June 1868, June 1879 (ny), Chickering s.n. (us), J. D. Smith s.n., oe Sept. 1884 (Gu, us), Shallert s.n., 1927 (puKE); Spruce Pine, Hyams s.n., June 1878 (us); Little Roan, Merriam s.n., | Sept. 1892 (us). Swain Co.: Great Smoky Mts. Natl. Park, Round Bottom Road { ea Heitooga Overlook, Athey s.n., 16 May 1973 (vps); 3.4 mi from N.C. hwy. 28 0 ica Knob Road, Bell 3117 (Ncu); Nantahala Gorge, 3 mi SE of Beechertown, Pa aan al. 8021 (NcU); Great Smoky Mts. Natl. Park, ets foot of Hughes Ridge, Jennison 445 (GSMNP, TENN); Great — Mts. Natl. Park, upper Chasteen Creek, Jennison & J. G. Smith 2291 (GSMNP, TENN); Brookside, Rich Mt., [unnewell 10337 (Gu). Transyly ania : Whitewater R. near Jct. a Bohaynee Road bridge, Bannister & Anderson 432 ‘u); near head of Davidson R. near Fish Rearing Sta., Pisgah speed Distr., Beaman 50026 (Nesc); 2.7 mi N of N.C. 215 on Courthouse Creek Road (U .S. Forest Service = — Map 1. Distribution of Diphylleia cymosa in the United States. [P86 VIATIAHdIC “TV LA ONIA 76 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Road 140), Boufford & E. W. Wood 16283 (Ncv), W Fork French Broad R., jet. U.S. 64 near Rosman, Bozeman et al. 9122 (BM, TENN); on trail, Black Rock to Sapphire, Coker s.n., 18 Aug. 1910 (NCU); Blue Ridge Pkwy. between Wagon Road Gap and Beech Gap, Freeman 57326 (Ncu); Pink Beds, House 4008 (Gu); road from Mt. Pisgah to Brevard, Oosting 34686 (pUKE); Pisgah Ridge, U.S. route 276 1.8 mi below ject. with Blue Ridge Pkwy., Roberts & Keil 7416 (vpn); Sapphire, NE-facing cove on Hogback Mt.. Ware & White 3055 (Ncu, vpB). Watauga Co.: 1.2 mi E of Bamboo on road to Triplett, Ahles & Ashworth 39619 (NCU), Blowing Rock, /. L. B. 3680 (puke), Rich Mt. NW of Boone, N-facing cove near lookout tower, Boufford & E. W. Wood 16310 (wcun), Chinese-Amer. Bot. ae 853 (KUN, NAS, PE, WH); Linville Road, Churchill s.n., 1899 (TENN); along Long Hope Creek on W slope of Old Field Bald, Hardin 13255 (Ncsc); upper end of Long Hope Creek, Hardin 13273 (Ncsc), Potato Hill, N of jet. of state roads 1324 and 1306, Kologiski & Perino 165 (GH, Ncsc); 2.5 mi SE of Aho, Radford 11111 (NCU), Rich Mt., Radford 45373 (Ncu, uscH); NW slope of Hanging Rock Ridge, Rohrer 1790 (Ncu), Grandfather Mt., Sevmour 91 8 13 (GH): 5 mi W of Blowing Rock, J. K. Small & Heller s.n., 1891 (G, Gu). Yancey Co.: 3.4 mi NW of Swiss, Ahles & Duke 50705 (Gu); Mt. Mitchell, Pa/mer ae (GH), Rhoades s.n., 1932 (Gu), Shallert s.n., 1923 (puUKE). SouTH CAROLINA. Oconee Co.: near $.C. route 107 near North Carolina line, Batson & Swails s.n., 1956 (scun); Ellicott Wilderness Area of Sumter Natl. Forest, E fork of Chatooga R., Kirkman & Ellis 1201 (Ga). TENNESSEE. Doe R. Valley, Ball s.n., Sept. 1884 (us). Blount Co.: Great Smoky Mts. Natl. Park, trail to Gregory’s, Cain s.n., 3 Aug. 1929 (TENN). Carter Co.: Roan Mt., | mi below Cold Spring, D. M. Brown 27 (pUKE); Roan Mt., below Carver’s Gap, Cannon 146 (Ny, Us): along old hacktrail leading to summit of Roan Mt., Wofford 81-36 (TENN). Cocke Co.: near Lemon’s Gap, Kearney 610 (NCU, NY, Us). Greene Co.: near summit of Cold Spring Knob, Sharp & D. K. Smith s.n., 23 Sept. 1973 (TENN). Monroe Co.: Cherokee Natl. Forest, Citico Creek WSA, Falls Branch Scenic Area, Malter 53107 (1eENN); Cherokee Natl. Forest, near Beech Gap, Sharp 715 (reNN). Sevier Co.: Great Smoky Mts. Natl. Park, Newfound Gap along U.S. route 441, Boom 39435 (1); Great Smoky Mts., “Spruce Flat,” Roaring Fork, Braun s.n., 25 April 1927 (us); Chimney Caps, Cain s.n., 12 June 1933 (TENN); Greenbrier, Lester Prong, Cain & Duncan 401:2 (TENN); Great Smoky Mts. Natl. Park, Chimney Caps Trail. Duncan 406 (Ga), Lester Prong, Duncan 90] (Ncu); Great Smoky Mts. Natl. Park, | mi E of Seilers Bald, Fosberg 18718 (Ncsc); Mt. LeConte, //tis 1355 (Ncsc); Smoky Mts., Balsam Point, ees ell ht (Gu); ee Pigeon R., Hunnewell 14178 (Gu); W fork of Little Pigeon R., ca. | mi W of Chimneys Campground, Hyypio 453 (vpp); Great Smoky Mts., Gr eee , near Ramsey Prone Creek, Jennison 39 (GSMNP, TENN): near Jakes Gap, Jennison 382 28 (GSMNpP); trail up Mt. LeConte, 7. Jones s.n., 4 May 1935 (1 om Mt. LeConte, Ramseur 1662 (NCU), el ee 2669 (TENN); Double Springs Gap. WW. B. Senopela 10038 (puke); Mt. LeConte, Roaring Fork, Sharp 640 (TENN): Great Lae Mts., Elkmont, Whmeyer 350 (GA). Unicoi Co.: E side of Rich Mt., valley of Higgins Creek, E. H. Cooley et al. 8855 (renn); Big Bald Mt., James s.n., 5 Sept. 1955 (reENN); along U.S. route 23, Flag ee James 16579 (TENN); Unaka Mt. near Beauty Spot, Price 702 (puke); near Bald Mt., Price 756 (UscH). VIRGINIA, Grayson Co.: Pine Mt., Solomon Branch, Sheffey s.n., 18 May 1974 (FARM); White Top Mt., Britton et al. s.n., 1892 (G, NY), Roller s.n., 1939 (vpi), Sharp 20496 (TENN), Stevens & Harvill 25604 (rarm), Uttal 11182 (vei); Mt. Rogers. Massey s.n., 1946 (vei), Nicely 826 (vPt), Reedy 65-4 (gency), C. E. Wood, Jr., 1403 (Gu, vp). Smyth Co.: White Top Mt., Camp 1564 (ny), Core (Moldenke) 6846 (ny), J. K. Small s.n., 28-29 May 1892 (GH, us), Stevens & Harvill 25656 (rArm); Mt. Rogers, Kral 1/684 (Ncu, vps). Washington Co.: Taylor’s Valley, Jervis s.n., 9 Aug. 1967 (EHCV). — — In addition to the two sheets of Diphylleia cymosa in the Michaux Herbarium at p (IDC 6211. 47: I. 2, 3), there are also two sheets in the general herbarium at p that are most likely part of the original collection and represent type 1984] YING ET AL., DIPHYLLEIA 77 material. Both of the latter sheets are annotated “Herb. Richard,” and both also bear labels with ““Herbarium Drake.” Background information on the role of L. C. Richard in the authorship of Flora Boreali-Americana and on the history of the Richard and Drake herbaria support this contention, as outlined below. Gray (1882a, p. 183), commenting on specimens in the “older herbaria,” said of Michaux’s Flora Boreali-Americana (1803), “It is known through tra- dition that this work was prepared by L. C. Richard, from the collections of the elder Michaux; but he wholly withheld his name, which therefore cannot be cited.’ Gray (1882b) also mentioned that Richard’s herbarium contained an almost complete set of the plants described in that work. Previous to Gray, Hooker (1842, p. 432) had written that “Richard 1s the anonymous author of the Flora Boreali-Americana of Michaux, in 2 vols. 1803.” It is of interest to note that Fedchenko (1937), in his treatment of the Berberidaceae for Flora SSSR, cited the authorship of Diphylleia as “L. C. Richard in A. Michaux.” Louis Claude Richard’s son, Achille Richard, continued his father’s botanical work and inherited his father’s herbarium (Gray, 1882b). The Richard her- barium later came into the hands of De Franqueville (Roze, 1891) and was eventually obtained by Drake del Castillo, who, according to Bureau (1904), acquired many of the larger private herbaria in Europe during the latter part of the 1800’s. In summarizing the contents of the Drake herbarium, Bureau mentioned that A. Michaux’s collections from North America were among them. Stafleu and Cowan (1976, p. 872) also mentioned the fact that the herbaria of L. C. and A. Richard are now at p through the De Franqueville and Drake herbaria. It appears obvious that the two specimens at p that are annotated “Herbarium Richard” are part of Michaux’s original collections of Diphylleia from North America and were among the material available to L. Richard when he prepared Flora Boreali-Americana. These two sheets, in addition to the two in the snes Herbarium, must therefore be considered in the designation of a lectoty The protologue of Diphylleia cymosa Michaux includes a description of all above-ground parts of the plant plus two plates (tt. /9, 20). One plate (t. 19) illustrates the habit of a plant in fruit (including a rhizome that more closely resembles that of Podophyllum peltatum L.); the other shows only the upper portion of a stem with the upper leaf and the inflorescence. Neither illustration exactly matches any of the four sheets of original material, although one sheet in the Michaux Herbarium (IDC 6211. 47: I. 2) and one in the general her- barium at Pp are quite similar to the illustration in ¢. 20 of Flora Boreali- Americana. The Michaux Herbarium specimen, however, 1s in fruit (as 1s [IDC 6211. 47: I. 3), while each of the two sheets in the general herbarium at p has both flowers and fruits. One of these sheets is labeled (in Michaux’s hand- writing?) with essentially the same information as in the original description. The characteristics of the attached specimens agree with the original description in Flora Boreali-Americana, and we designate the flowering material on this sheet as the lectotype of D. cymosa Michaux. Although the three species of Diphy/leia are remarkably alike. D. cymosa appears to be most similar in overall morphological features to D. sinensis, of 78 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 central and southwestern China. Both have a petiolate, peltate upper leaf, smaller flowers, more abundantly flowered inflorescences, and tricolpate or hexarugulate pollen—characters not found in D. grayi, of Japan. The inflores- cences are commonly cymose in D. cymosa and D. sinensis but are frequently umbellate in D. grayi. However, D. sinensis and D. grayi are similar in the pubescent inflorescence, the greater number of ovules per ovary, and the exine ornamentation of the pollen. It is difficult, if not impossible, to say with cer- tainty which species is most primitive. 2. Diphylleia sinensis H. L. Li, J. Arnold Arbor. 28: 442. 1947. Figure 14. Diphylleia cymosa Michaux subsp. sinensis (H. L. Li) T. Shimizu, Hikobia Suppl. 1 450. 1981. Plants 4-10 dm tall, growing in dense to loose co lonies or individually. Petiole of lower leaf 7-20 cm long, of upper (2.5—)6-13 cm long; blade peltate, reniform or reniform-orbicular to transversely oblong, 19-40 by 20-46 cm in lower leaf and 6.5-31 by 19-42 cm in upper, 2-cleft with divisions undulate or shallowly 3- to 6-lobed, margin irregularly dentate with teeth apiculate, pubescent with whitish hairs beneath, sparsely so or subglabrous above. Peduncle 3.5-28 cm long, solitary, occasionally bifurcate; inflorescence 4.2-35 cm long including peduncle, 3.5-10 cm broad, branches pubescent or glabrescent in fruit. Flowers (8 to) 15 to 51; pedicels 0.4-3.7 cm long; outer sepals lanceolate to linear- lanceolate, 2.3-3.5 A 0.7-1.2 mm: inner sepals very broadly elliptic to sub- ee 4-4.5 by 3.8-4 mm, outer petals narrowly to broadly obovate, 5-8 by 2.5-5 mm; inner petals narrowly elliptic to narrowly obovate, 5.5—-8 by 2.5- ay mm: stamens ca. 4 mm long, laminar filaments 1.7—2 mm long, anthers ca. 2 by 0.6 mm: ovary ellipsoid, 3-4 by 1.8—2 mm, ovules 5 to 11, style absent or represented by slight constriction at summit of ovary, stigma ca. 0.3 by |! mm. Berry globular to broadly ellipsoid, 10-15 by 6-10 mm broad, dark blue or purple-black, on pedicel 1-3 cm long. Seeds 2 or 3, ovoid, ca. 5.5 by 3.5 mm, + rounded on all sides or only slightly flattened ventrally. Chromosome number: unknown. Type. China, western Sichuan Province, July & August 1908, £. H. Wilson 814 (lectotype, GH (the fruiting element); 1solectotype, Us). DISTRIBUTION AND ECOLOGY. Moist deciduous and coniferous forests, some- times bamboo thickets, 1880-3700 m alt. China: western Hubei, southern Shaanxi, southern Gansu, Sichuan, and northwestern Yunnan. (Map 2.) SPECIMENS pate Chi nina. GANSU: Zhouchu Xian, P. Z. Guo 5148, 5567 (wuk), S. Zhang 00406 (PE), = Zhou 294 (nwrc), Zhang Xian. Y. S. Lian 790160 (xwrc); Dang Chang X1an, : . Lian 790971 (nwrc); Die Bu Xian, ¥. S. Lian 800328 (Nwic C. R. Wang 15643 oe Nc); Liou hia tien ze, col du Koan Chan, /. Licent 5325 (3m, kK, p), 5335 (kK), 5336 (kK); Tianshui Xian, J. M. Liu 10182 (ee), Z. W. Zhang 151 (wux); Ming Xian, J. C. Wang ee VYu-Mo-Tin Team 387 (wnc), Longde Xian, Z. P. Wang 13120 (wu); Xigu Xian, Z. P. Wang 15159 (pe, wuk); Jing Yuan, 7. P. Wang 17074 (pe); Wu Shan Xian, ne collector 324 (nwrc). Huser Badong Xian, Ff. /7/. — 1984] YING ET AL., DIPHYLLEIA 79 pa PAN AR wy Hi? f ij Ee oy ae oy, s A NY yy | Figure 14. Diphylleia sinensis (based on Nan-shui-Bei-diao Team 7164, G. X. Fu 1062, Shennongjia Bot. Exped. 10276: all pe): a, upper portion of plant, rhizome. roots: b, flower: c, outer sepal: d. inner sepal; e, outer petal: f, inner petal: g. stamen; h, ovary: i, apical bud of young plant (from top of rhizome); j, undersurface of leaf. 80 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Chen 5132 (nip), G. X. Fu 1062 (pe), R. H. Wang 307 (nip), Fang, 4. Henry 6820 (3M, E.G, GH, K, NY, US), E. H. Wilson s.n. (w); Shennongjia Forest Distr., Shennongyia Bot. Exped. 10276, 22868, 25107, 31388, 32451 (Hib, PE), Shennongjia Forest Distr., X1ao- shennongjia, Shennongjia Bot. Exped. 10599 (a, Hi, pe), Shennongjia Forest Distr., vic. of Dalongtan and Xiaolongtan, /980 Sino-Amer. Bot. Exped. 910 (A, HB); ““W Hupeh,” BE. H. Wilson 2112 (w). a ee Mei Xian, sae Shaanxi ae 85 (wuk), Z. P Wang 19976 (wuk), Feng Xian, K. J. Fu 8306, 12299, 13285 (wuk); pee Shan, K. J. Fu 4452, 9424 (wur), P. y. i 5199 (wuk), } vs oe 125 an K), 7. N. Liou & P. C. Tsoong 330 (ve, WUK), 2263 (PE), Qinling Team 300 (wuk), C. L. ae ee (WUK), Z. P. Wang 15741 (wur), G. P. Wei 1492 (wuk); Gua-in-san, G. Giraldi 1867 (F1), Kin- tou-san, G. Giraldi 6004 (F1); Thae-pei-san, e Giraldi 6274 (1); Hu Xian, P. Z. Guo 386 (KUN, PE); Fuping Xian, Daping, B. G. Guo 1421 (pr, wuK), 1769 (A, PE); Weinan, Z. P. Wang 15632 (pr). SICHUAN: Tchen-keou Xian, 7. R. Dar eee Ore R. P. Farges 58P (« (2 sheets), p (3 sheets)). Kangding Xian, B. C. Gao et al. 55 (sz); Heishui Xian, San-dao-gou, Heishui Team 1075 (A), 1354 (pr), X. Li 73212 mae (pe); Li Xian, Z. Ho 13376 (ve), Z. R. Wu 33348 (re); Jinfu Shan, Z. Y. Lau 780390 (rr), G. FF. Li 62274 (pe), Maowen Xian, Maowen Team 2642 (pe); Heishui Xian, Nan- shui- Bei- diao Team 01525 (re); Mealo ae pe -su-gou, Nan-shui-Bei-diao Team 7164 (A, PE). ““W Szechuan” [Wenchuan Xian, near Wasi], E. H. Wilson 814 eg K, US). YUNNAN: Ma- cul-chan, Delavay 3862 (A, P O sheets)3), s.m. (kK); “Yun-nan,” Delavay s.n. (A, P). G. Forrest 30037 (3m, ©); Chungtien mae Feng 1457 (KUN); Pa Xian, Feng 6382 KUN, PE); Lichiang Range, NW flank, G. Forrest 10488 (Bm, ©, L); Chungtien Plateau, G. Forrest 12524 (Bm, £); “duplicate ’ 1913-1914,” G. Forrest 15458 (e, «); “duplicate of 1917.” G. Forrest 18709 (©, Us); Landsang-djiang (Mekong) & Lu-djiang (Salween). vic. of Tseku. H. F. Handel-Mazzetti 8921 («, w (2 sheets)); Lichiang Xian, Lichiang Bot. aie 100267 (rE); Mt. Fu-Chuan, H. D. McLaren’s coll. 203 (&), Lichiang Range, H. McLaren’s coll. 203d (pm), Yangtse watershed, Lichiang Snow Range, J. fF. Rock re us); Mt. Fu-Chuan, SW of Wei-Hsi, Mekong-Salween, J. F. Rock 16971 (&, GH, NY, US) The type of Diphylleia sinensis (Wilson 814, Gu) is based on specimens— one in flower and one in fruit—collected at different times of the year and mounted on the same sheet. The single label bears two dates, “7/08 + 8/08.” Li (1947) did not indicate a choice of elements in designating the type, but in 1951 he annotated as “isotype” a duplicate of Wilson 814 (us) consisting only of fruiting material. Since flowering and fruiting material are about equally represented on the GH specimen and each applies equally well to the original description, we have chosen the fruiting material as the lectotype. Although no locality more exact than “Western Szechuan” is given on the label of Wilson 814, it is probable that Wilson’s collections of Diphylleia sinensis came from the area in Wenchuan Xian near Wasi (““Wa-ssu”’) at about 31°28'N, 103°28’E (see Clausen & Hu, 1980; Howard, 1980). This is based on the fact that Wilson’s numbers 8/3 and 8/5 (Rubus pileatus Focke and R. giraldianus Focke) were both collected at that locality (Rehder, 1913; Sargent, 1916) 2Farges 581 ranges from plants in bud to plants with mature fruits, the specimens were obviously collected over several months and later lumped under a single numbe ‘The specimen at a is in flower, while the 2 specimens at p are in fruit. The date on the labels is 6 August 1889, obviously an error on the a sheet. 1984] YING ET AL., DIPHYLLEIA Distribution of Diphylleia sinensis in China. Map 2. 82 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 3. Diphylleia grayi F. Schmidt, Mem. Acad. Imp. Sci. Saint Pétersbourg 12(2): 109. 1868. (Reis. Amurl. 109. 1868). FIGURE 15. Diphvileia grayi F. Schmidt var. (ypica H. Takeda in H. Takeda & K. Tanabe, Kozan Shokobutsu Shashin-jushu, 95. figs. 134-136. 1931. Diphylleia grayi F. Schmidt var. incisa H. Takeda in H. Takeda & K. Tanabe, Kozan Shokobutsu Shashin-jushu, 96. figs. 137-139. 1931. Type: no specimens were cited in the original description, and we have seen no specimens collected or annotated by Takeda as var. incisa. In lieu of a specimen, we designate fig. 139 in Takeda and Tanabe (1931) as the lectotype. Diphylleia grayi F. Schmidt var. rotundata H. Takeda in H. Takeda & K. Tanabe, Kozan Shokobutsu Shashin-jushu, 97. Lae Me 1931. Type: no specimens were cited in the original description, and we have seen no specimens labeled as var. rotundata by Takeda. In the absence ofa specimen, we ase /40 in Takeda and Tanabe (1931) as the lectotype. Diphylleia cvmosa Michaux subsp. grayi Be hmidt) Kitamura in Kitamura & Mu- rata, Acta Phytotax. Geobot. 20: 202. 1962 Diphylleia cvmosa Michaux subsp. grayi (F. ne Kitamura var. incisa(H. Takeda) T. Shimizu, Hikobia Suppl. 1: 450. 1981. Plants 2.5-9 dm tall, the stems with scattered, curled, white pubescence; growing in dense to loose colonies or sometimes individually. Lower leaf with petiole 1-25 cm long, blade 9.2—35 by 11-44 cm; upper leaf sessile or occa- sionally with short petiole to | cm long (very rarely up to 5.8 cm long), blade attached at sinus, not peltate or only slightly so, 3.9-27 by 4.8-35 cm; blades of both leaves orbicular to reniform-orbicular, margin doubly serrate and sometimes deeply lobed, pubescent pense staal SO oe Peduncle 0.8- 7cm long, to 16 cm long in fruit, pubescent, som s densely so: inflorescence 2.2-21 cm long including peduncle, 2-8 cm broad, branches nce Flowers (2 to) 4 to 16, on pedicels 0.8-3.2 cm long; outer sepals lanceolate to linear- lanceolate, 4-7.2 by 0.6-1.2 mm: inner sepals broadly elliptic, 4-9 by 3-5.4 mm, outer petals broadly obovate, 8-15 by 6-11 mm, inner petals broadly obovate, 8-15 by 4.8-11 mm; stamens ca. 4 mm long, filaments ca. 0.7-1.5 mm long, anthers ca. 2—3.2 by 0.8-1.4 mm; ovary ellipsoid, 2.5-5 by 1.7-2.8 mm, ovules 5 to 11, style 0.3-0.8 mm tall, stigma 0.2-0.7 by 0.8-1.5 mm. Berry ellipsoid to subglobose, 9-18 by 8-16 mm, blue, on pedicel 0.9-4 cm long. Seeds (3 to) 5 to 7 (to 10), ovoid to oblong, ca. 6-6.5 by 3.3-4.5 mm, rounded on all sides. Chromosome number: 7 = 6 Type. Schmidt (1868) cited the following specimens (all from Sakhalin), which we have not seen, in his description of Diphylleia grayi: Arkai, 27 May 1861, Glehn s.n., beginning of August 1860, Glehn s.n.; Dui, beginning of June 1860, and 21 July, Glehn s.n., Estaing Bay, July, Bry/kin s.n., Kussunai, 25 April 1861 and mid-May, Brylkin s.n., Manue, mid-August 1860, without ceaaal or number. All are syntypes and are presumably at LE, where Gle and Brylkin’s specimens are deposited (Holmgren, Keuken, & Schof aL on Lanjouw & Stafleu, 1957; Stafleu & Cowan, 1976). DISTRIBUTION AND ECOLOGY. Moist places in cool deciduous and coniferous forests, usually along small streams and seepages: from near sea level in the 1984] YING ET AL., DIPHYLLEIA 83 Figure 15. Diphylleia grayi (based on Brooks 66/, pr): a, upper portion of plant. rhizome, roots: b, flower: c, outer petal: d. inner petal: e, stamen: f, ovary. 84 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 north to 2700 m in the south. U.S.S.R. (Sakhalin, below 50°N), Japan (from Hokkaido (throughout) to southwestern Honshu (mostly limited to western side)), Kuril Islands (Kunashir1). (MAP 3.) = CIMENS EXAMINED. Japan, HOKKAIDO (Hokkaido Prefecture). “Southern Hokkaido,” ’ D. Brooks 661 (ee). Abashiri Shicho: Shirataki-mura, Mt. Taira-yama, K. /fo s.n. i Ap): Shari-cho, Raus-daira, S. Kobayashi s.n., 20 July 1967 (mak); Shari-cho, Mt. Shari- dake, S. Kobayashi s.n., 31 July 1967 (mak); Takinoue-cho, S. Okamoto s.n., 22 July 1952 (kyo); Mombetsu city, Shokotsu, S. Okamoto s.n., 5 Aug. 1954 (kyo). Hidaka Shicho: Hiratori-cho, Shoya, K. Jto s.n., 11 May 1963 (sap); aap Mt. Petekari- dake, H. Koyama & N. Fukuoka 3256 (1ns); Hiratori-cho, . Horoshiri-dake, //. Koyama et al. 3591 (tNs); Samani-cho, MM. Tatewaki 9242 ee Erimo-cho, Saruru- sando, M. Tatewaki 42844 (SAP), . Tokubuchi s.n., 14 Aug. 1892 (Ny, sap). Hiyama Shicho: Okushiri-cho, Okushiri Is., K. Mivabe & Tokubuchi s.n., 28 July 1890 (k, sap), K. Togashi 21821 (sap). Iburi ae Muroran city, Mt. Washibetsu, S. Kawano 444 (sAp); Shiraoi-cho, Mt. Orofure, S. Kawano 530 (sap). Ishikari Shicho: Sapporo city, Mt. Soranuma-dake, S. Adivama s.n., 19-20 June 1932 (ny), H. Hara s.n., 18 June 1937 (TNs), Y. Yokomizo s.n., 22 June 1958 (sap); vic. of Sapporo, S. Arimoto s.n., 26 May 1903 (GH, MoO); Sapporo city, Jozankei, S. Kurosawa & Y. Tateishi s.n., 2 June 1975 (KYO, Tl, TUS), J. Ohwi 4662 (Kyo), S. Terabayvashi S88 (Kyo); Eniwa city, Mt. Eniwa, Mivabe s.n., 7 Aug. 1902 (sap): Sapporo city, Mt. Teine-yama, S. Saito 456 (Gu), H. Yanagishima g.m., 1 June 1912 (sap); Sapporo city, H. Takeda s.n., 13 May 1909 (BM, E, K, US); Moiwa-yama, H. pee sn, 6 June 1923 (sap), alegre Shicho: Kamikawa- oe Mt. Taisetsu-san, N. Kitagawa 1153 (Kyo), T. Nakai s.n., Aug. 1928 (rt), J. Ohwis.n., 9 Sept. 1928 (TNs), 5 Okuyama s.n., 18 July 1941 ee Minamifurano- cho, Mt. Yubari-dake, G. Murata & 7. Yahara 37461 (Kyo, sap); Otoineppu-mura, G. Murata et al. 38447 (kyo); Furano city, Furano, S. Nishida s.n., 25 July 1916 (sap); —~ Kamikawa-cho, Mt. Yul-ishikari-dake, J. Samejima & T. ale a et) Kushiro Shicho: een cho, Mt. Mashu-dake, K. Takita 297 (kyo). M : Nemuro city, Nosappu, H. Toda s.n., 15 July 1954 (rus). Oshima Shicho: Ono-cho, Naruka Valley, F. cE Cu 236 (sap); Matsumae-cho, Mt. Daisengen-dake, 7. Ishivama (sap); Kamuso-cho, Moheji, S. ee sn, 25 May 1906 (sap); Hakodate. C. Maximowicz s.n., 1861 (BM, FI, G, GH, K, L, NY, US); Assabu-mura, Uzura R., 7. ee & Maruyama s.n., 22 May 1950 a. Nanae-cho, Mt. es Nek H. Takeda s.n., 21 July 1977 (Kyo, TI), /. Yamamoto 566 (sar); Fukushima-cho, Mt. Daisengen-dake, K. Takeda s.n., 24 July 1977 (Ncw); Nanae- ra co ma, A/, eee 44602 (SAP); Yakumo-cho, Kiootoshike, M. Tatewaki et al. s.n., 25 May 1955 (sap); Oshamanbe- cho, Inaho Pass, Y. Tokubuchi s.n., 16 July 888 es Rumoi Shicho: Horonobe-cho, beageae H. Koyama 1688 (kyo), Horonobe-cho, Mt. Chikoma-dake, V/. ions , 12 July 1931 (sap); Mashike-cho, Mt. Shokanbetsu-dake, /. Yamamoto 5474, 5 ae Shiribeshi Shicho: Otaru, mountain gorges, U. Faurie 201 (vp), Takenouchi s.n., 38 May 1928 (L); Niki-cho, Nakanosawa, S. Ishikawa s.n., 2 July 1893 (sap); Mt. Tengu- yama, Kawata s.n., 12 May 1935 (sap); Shimamaki-mura, along Tomari R., 7. sea & J. Igarashi s.n., 28 June 1954 (sap): Iwaoto, B. Yoshimura & H. Yokoyama s.n., June 1938 (sap). Sorachi Shicho: Ashibetsu city, Mt. Kirigishi-yama, /. /shizuka & 7 Tohyama s.n., 26 July 1970 (sap); Ashibetsu city, Mt. Ashibetsu-dake, /7. Koidzumi 67192 (tNs); Uryu- cho, Mt. Minamishokan-dake, G. Murata et al. 38644 (CM, KYO, SAP), 38695 (KYO, SAP); Yubari city. Mt. Y ubari-dake, : ae 5041 (Kyo), K. Sugawara s.n., 13 July 1969 (rus); Yubari, } eee i SM, . 1892 (Gu). Soya Shicho: Nak- atonbetsu-cho, Af. Yamanoi s.n., 28 May 1972 ie — ax Soya, C. Wright s.n., 15 July 1853-1856 (Gu, Ny). Tokachi Shicho: Shikaoi- -mura, near Lake Shikaribetsu, MM. Wak- abayvashi 738 (kyo). Honsnu. Mirosaki, U. Faurie 574 (wu). Akita Prefecture: Senboku- gun, Tazawako-machi, Hachimantal, /f. Hara s.n., 20 July 1953 (11), S. Kurosawa s.n., 4 Aug. 1951 (11); Senboku-gun, Tazawako-machi, Mt. Komagatake, HW. Hara s.n., 8 July > Oo 1984] YING ET AL., DIPHYLLEIA are f Jo¥ ~ ( a a Nass eo sg “eer? ee °. ny .* oe 300 km ~ Map 3. Distribution of Diphylleia grayi in Japan and the specimens; triangles bas _S.S. R. Dots based on ed on literature reports (Board eee of Iwate Prefecture, 1970; Muramatsu, 1932; 2: M. Noda, 1969; Sugawara, 1937). 86 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 1958 (ri), HT. Muramatsu s.n., 7 Aug. 1928 (11); Yuri-gun, Chokai-machi, Mt. Chokai T. Kaneko s.n., 15 Aug. 1962 (rus); Kitaakita-gun, Ohta-machi, Mt. Yakushi, Kurata sm, 3 June 1978 ‘ge Pref. Mus.); Kazuno-gun, Kosaka-machi, Mt. Shirozi-yama, R. Mochizuki s.n., 20 July 1969 (herb. Mochizuki); Hiraka-gun, Sannai-mura, Nango, R Mochizuki s.n., 3 May 1977 (herb. Mochizuki); Yuri-gun, Ouchi-machi, Shirouchi, R. Mochizuki s.n., 7 May 1977 (herb. Mochizuki); Honjo city, Kitanomata, R. Mochizuki s.m., 11 May 1980 (herb. Mochizuki); Senboku-gun, Nishisenboku-machi, Osawago, R. Mochizuki s.n., 6 June 1981 (est: Mochizuki); Kitaakita-gun, Tashiro-machi, Mt. Tash- iro-dake, R. Orkawa s.n., 25 May 1968 (rns); Kitaakita-gun, Ohta-machi, Kitamahiru, Takada s.n., 7 June 1978 (Akita Pref. Mus.); Ogachi-gun, Minase-mura, Mt. Toraga- yama, Tohoku Univ. Bot. Gard. Staff s.n., 23 July 1970 (mo, rus); Ogachi-gun, Ugo- machi, Karuizawa, Tsuchida s.n., 10 May 1974 (Akita Pref. Mus.); Ogachi-gun, Minase- mura, Mt. Kiji-san, 7’suchida s.n., 10 May 1974 (Akita Pref. Mus.); Akita city, Mt. Ohira-yama, Yoshikawa s.n., 15 June 1955 (Akita Pref. Mus.). Aomori Prefecture: Aomo- ni, U. Faurie 574 (Bm); Hakkoda, U. Faurie 926 (F1), 928 (G), 13072 (mo, p); Aomori city, Hakkoda Mts., Mt. Odake, 7. Fiyita & E. Nakagawa s.n., 1 July 1908 (tus), A. Kimura & S. Sugaya s.n., | July 1953 (rus); Kamikita-gun, Towadako-machi, Hakkoda Mts., Sarukura, K. Hasegawa s.n., 30 June 1964 (11), S. Sugaya et al. s.n., 2 July 1955 (rus); Kamikita-gun, Noheji-machi, Mt. Eboshi-dake, K. Hosis.n., 17 June 1951 (KANA); Shimokita-gun, Sai-mura, O. Mori 37 (MAK), Kamikita-gun, Towadako-machi, Mt. To- wada-yama, 7. Naito 7750 (rus); Minamitsugaru-gun, Hiraga-machi, Mt. Kushigamine, Osawa Senior High School aeons oe Nakatsugaru-gun, Le machi, Mt. Iwaki- san, S. Narita s.n., 19 June 1910 (MAK), H. Sakurai ae Sept. 1885 (TNs); Kamikita- gun, Tohoku-mura, Kochi. Z. ashing SM, July 1948 a Fukui Prefecture: Katsuyama city, Tani-toge Pass, G. Koidzumi s.n., 7 He 1921 (Kyo): Ono city, ML. Beko-san, 2. Tashiro s.n., = June 1937 (TNs); Nanjo-gun, Imajo-machi, Yashagaike Pond, N. Fukuoka 4889 (kyo), K. Ueda S08 (A, KYO, MO, T1); Katsuyama city, Mt. Tottate-yama, S. ae ae s.4., 16 May 1964 (kyo). Fukushima Prefecture: Yama-gun, Inawashiro-machi, Mt. Azuma-yama, G. ee s.n., June 1904 (rns), S. Okuwvama — 3558 (TNs); Minamiazumi-gun, Hinoemata-mura, Mt. Hiuchi, G. Nakai 2587 (Kyo): Minamiazumi-gun, Tateiwa-mura, Mt. Tashiro-y: yama, S. ede ee (TNS); Mina- miazumi-gun, Tadami-machi, Mt. Asakusa-dake, Mf. Suzuki s.n., 12 July 1960 (rNs); Nishishirakawa-gun, Nishisata-mura, Mt. Asahi-dake, /. a sn, 29 July 1958 (1Ns); Onuma-gun, Showa-mura, Komado, without collector or number, 10 May 1905 (MAK). Gifu Prefecture: Ibi-gun, Sakauchi-mura, SW of Mt. Sobatsubu-yama, N. Fukuoka S793 (KANA, KYO); Ono-gun, Shirakawa-mura, Hirase, VW. Hashimoto 382 (KANA); Ono- gun, Shokawa-mura, Tenbu Pass, J. Hatasa s.n., 25 July 1968 (Ns); Ibi-gun, Kasuga- mura, E of Mt. [buki- Po M. Hutch 23487 (Kyo), Y. Satake & S. Okuyama s.n., 24 May 1942 (1Ns); Ono-gun, Takane-mura. Nomugi Pass, H. Kanai s.n., 13 June 1958 (MAK), Hf. Kanai & H. "Ohashi 731217 (rns); [bi-gun, Sakauchi-mura, Kawakami, //. Kanai & T. Morita 0259 (11); Masuda- sae Kasako-cho, NW of Mt. Ontake-san, G. Murata 10670 (kyo); Yoshiki-gun, Mannami, Y. Nagai s.n., 31 May 1972 (KANA, MAK), N. Naruhashi 3616 (KANA), Gujo-gun, ene mura, _ Daihi-dake, S. Ueno SM, 22 July 1970 (rns). Gunma Prefecture: Shimizu-toge Pass, U. Faurie 2396 (mo); Tone- gun, Kaashina-mura, Oze, 7. Harazawa s.n., 1 Aug. 1910 (MAK), Af. Mizushima 1121 (11), G. Nakai 2628 (Kyo), . Okuyama s.n., 21 mas 1934 ( M. Ono s.n., 2 June 1954 (11); Tone-gun, Minakami-machi, ML. "Tanigaw M. Furuse s.n., 16 June 1958, 6 June 1961 (A), K. Hasegawa s.n., 15 July 1966 (71), K Hisauchi 489 (4), H. Kanai 3368 ri), 7. Miyamae s.n., 30 May 1969 (rns), R. Noguchi s.n., 24 June 1934 (rns), S. mee T3055 (tws), 7. Yamaz akt g.a., 24 June 1944 (11); Tone-gun, Minakami- achi, Mt. Hotaka, H. Kanai s.n., 5 July 1957 (11); Tone-gun, Minakami- si Mt. Hirapa- take, f/. Kanai s.n., 1 Aug. 1959 (rt): Azuma-gun, Himekoi-mura, Hiramata, K. Matsuda 378 (1ns); Tone-gun, Kaashina-mura, Mt. Shirane, H!. Ohba & S. ae 317 (A, Kyo, TH; Tone-gun, Nitharu-mura., Mikuni Pass. S. Okuvama et al. s.n., 11 ae 1984] YING ET AL., DIPHYLLEIA 87 July 1957 (rns). Hiroshima Prefecture: Hiba-gun, Mt. Sarumasa-yama, G. Akivama s.n., 23 May 1976 (1Ns); Mt. Omaki-san, 5. Takafuji 583 (kyo). Hyogo Prefecture: Kinosaki- gun, Hidaka-cho, Mt. Sofugatake, E. Araki 3123 (Kyo); Mikata-gun, Muraoka-cho, Mt. Torokawa-yama, E. Araki s.n., 12 July 1931, 31 August 1931 (kyo), M. Hashimoto 9223 (kyo), G. Koidzumi s.n., July 1927 (Kyo), G. Murata 20902 (Kyo), Mikata-gun, Onsen-cho, Mt. Ogino-sen, S. Hosomi s.n., 31 July 1968, 4 May 1969 (Kyo); Yabu-gun, Sekinomiya-cho, Mt. Hyono-sen, G. Murata & K. Iwatsuki 700 (Kyo, TNS), J. OhWi S.n., 6 July 1927 (1Ns), S. Terabayashi 60 (Kyo), Y. Yoneda s.n., 13 May 1933 (Kyo). Ishikawa Prefecture: Kanazawa city, Komagaeri-Kurotani, Mt. Takasaburo-yama, N. Fukuoka 7658 (KYO); Kanazawa city, Mt. Takao, /. Kaneko s.n., 20 May 1962 (KANA, MAR); Ishikawa-gun, Shiramine-mura, Mt. Haku-san, U. Faurie? 2798 (pe), M. Hotta 5707, 1853-1856, (kK); Kamiminochi-gun, Kinasa-mura, Okususobana, S. /to 152 (17Ns), Min- amiazumi-gun, Azumi-mura, Mt. Norikura-dake, 7. Itos.n., 19 Aug. 1891 (11); Kamiina- gun, Hase-mura, Mt. Senjo-dake, K. Iwatsuki et al. 120 (kyo), G. Murata 8275 (Kyo), T. Yamazaki s.n., 3 Aug. 1949 (11); Kiso-gun, Oshika-mura, Mt. Akaishi-dake-Mt. Osawa-dake, K. Iwatsuki & H. Koyama 183 (kyo); Shimoina-gun, Achi-mura, Mt. Ena- san, H. Kanai s.n., 29 July 1957 (11); Shimotakai-gun, Yamanouchi-cho, Shigakogen, H. Kanai 7607 (11); Shimoina-gun, Oshika-mura, Ogawara, 7. Kawamata 65 (1Ns): Minamiazumi-gun, Azumi-mura, Tokugo Pass, G. Koidzumi s.n., 31 May 1928 (kyo). T. Nakai s.n., 11 July 1927 (11); Minamiazumi-gun, Azumi-mura, Kamikochi, A. Kojima s.n., 6 July 1934 (Kyo), K. Ueda et al. 19 (Kyo); Kiso-gun, Oshika-mura, Sanpuku Pass, F. Konta 10326 (kyo), J. Sugimoto 4915 (11); Nishikomaga-dake, M. Kume 2595 (a); Kitaazumi-gun, Hakuba-mura, Mt. Shirouma, 7. Makino s.n., Aug. 1918 (MAK), N. Maruyama s.n., 2 Aug. 1950 (rns), M@. Sugivama $.n., 29 June 1968 (11), S. Terabayashi 306, 793 (KYO), F. Tomihisa s.n., 16 Aug. 1923 (Kyo), Y. Yabe s.n., 26 Aug. 1902 (11); Kitaazumi-gun, Hakujo-mura, to Sarukura from Futamata, foot of Mt. Shirouma, AV. Furuse s.n., 25 May 1958 (a); Kiso-gun, Minamishinano-mura, Mt. Hiuiri-dake, /7. Matsuda s.n., 17 June 1954 (11); Omachi city, Taira-mura, 7. Mimoro et al. 1492 (TNs): Shimotakai-gun, Kijimadaira-mura, Kayanodaira, M. Mizushima s.n., 16 June 1966 (A, KYO, MAK, TI); Kiso-gun, Oshika-mura, Mt. Otaka-san, M. Muramatsu 1487 (vs); Ki- 88 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 taazumi-gun, Otari-mura, Mt. Amakazari, D. FE. Boufford & E. W. Wood 19145 (a), G. Murata et al. 30378 (kyo), S. Terabayashi 767 (Kyo), S. Tsugaru 3389 (mo); Minami- azumi-gun, Hotaka-mura, Mt. Tsubakuro, ¥. Ogura s.n., 25 June 1918 (11), Y. Satake s.n., 13 July 1941 (rns); Kamiina-gun, Hase-mura, Mt. Kitaarakawa-dake, Y. Okada s.n., 12 Aug. 1923 (1Ns); Chino city, Mt. Yatsuga-take, C. Okawa s.n., 4 July 1977 (Ns), Y. Yabe s.n., 20 Aug. 1902 (11); Minamiazumi-gun, fader a Mt. Otaki, S. Okuvama etal. s.n., 18 Aug. 1954 (rns); Kamiina-gun, Hase-mura, Mt. Komaga-take, H. Sakurai g.m., 10 Sept. 1884 (rns); Kiso-gun, Oshika-mura, Mt. Toyopuchi. -san, 7. Yamazaki s.n., 29 July 1953 (11); Kiso-gun, Agematsu-mura, Mt. Kisokoma-dake, without collector or nuraber, July 1935 (MAK). Nara Prefecture: Mt. Sanjogatake-Mt. ee Gyo- jagaeri, Hf. Hara ie 15 July 1955 (m1), M. Hotta et al. 55 (kyo), G. M ad& K. apie 72 (KYO), ¥. Momivama s.n., 16 July 1955 (71); Yoshino- pin. Te se awa-mura, Omine Mts., Mt. Misen-Gyojagaeri, 7. Hotta et al. 74 (Kyo), M. Hotta & N. Fukuoka 258 (KYO), G. Koidzumi s.n., 14 July 1922 (kyo), G. Murata & T. Shimizu SS (KYO), H. Okada et al. 1370 (Kyo), S. Tanaka s.n., 17 July 1933 (k, TNs), Yokohama Nursery Co. s.n., Aug. 1907 (k). Niigata Prefecture: Nishikanbara-gun, Yahiko-mura, Mt. Yahiko, N. Fukuoka 1828 (kyo); Nishikubiki-gun, Myokokogen-cho, Mt. Kurohime-yama, NV. Fu- kuoka 2158 (Kyo); Nishikubiki-gun, Myokokogen-cho, Mt. Myoko-san, 8. Matsuda s.n., 27 July 1894 (Kyo), G. Murata 6834 (kyo), Nishikubiki-gun, Myokokogen-cho, Sasa- gamine, A. Nitta 106817 (Kyo); Nakauonuma-gun, Tsunami-mura, Mt. Naeba, S. Oku- yvama s.n., 19 July 1936 (Ns), Y. Satake & H. Ito s.n., 12 July 1951 (tNs), Shiga Prefecture: Higashiasai-gun, Asai-cho, Takayama-Torigoe Pass, Y. Jnamasu & N. Fu- kuoka 82 (kyo); Ika-gun, Suino-mura, Mt. Tsuchigura, Y. Inamasu & N. Fukuoka 205 (KANA, KYO); Higashiasai-gun, Ibuki- a ea Ibuki, G. Koidzumi s.n., 26-27 June ne (KYO); pane -gun, Asai-cho, Mt. Kanakuso, G. Murata & N. Fukuoka 1020 (ke, , MAK, MO, TNS); ae ore [mazu-cho, Mt. Miedake, 5S. Watanabe 5.1, 26 ae 1927 (KYO). Shimane Prefecture: [ishi-gun, Yoshida-mura, Mt. Omaki-san. Moriyama s.n., 5 Aug. 1961 (MAK). Shizuoka Prefecture: Shizuoka city, Mt. Hakkorei, . Konta et al. 309 (Kyo); Shizuoka city, Mt. Senmai, H. Matsuda s.n., 7 July 1954 (1); Shizuoka city, Higashimata, H. Matsuda s.n., 22 July 1954 (11). Tochigi Prefecture: Nikko city, Mt. Shirane-yama, J. Bisset 4027 (re), Hattoris.n., 1 July 1922 (11), 7. Makino sn, 17 July 1924 (mak), H. Sakurai s.n., 16 July 1887 (rns); Shioya-gun, Fujiwara , kahara-yama, H. Kanai s.n., 25 May 1959 (11); Nikko city, Yumoto, S. Kobayashi 45870 (MAK), G. Murata 18205 (KYO), N. Shibusa s.n., 2 July 1966 (tNs), S. Suzuki s.n., 21 June 1931 (Kyo); Nikko, Yokohama Nursery Co. s.n., Aug. 1906 (e). Tottori Prefecture: Saihaku-gun, Daisen-cho, Mt. Daisen, S. Horis.n., 4 Aug. 1889 (MAK), N. Kinashi s.n., 7 June 1917 (Kyo), G. Koidzumi s.n., 3 July 1924 (kyo); Yazu-gun, ae mura, Kitadani, G. Murata et al. 298 (Kyo). Toyama Prefecture: Shimoshinkawa- -gun Aeahi- cho, Mt. Iburi, #7. Kanai s.n., | Aug. 1958 (11); Higashitonami-gun, Minoya-mura, H. Kaneko s.n., 19 May 1963 (kKANA, 11); Nakashinkawa-gun, Paine ene cho, Takamine-yama, N. Kurosaki 16017 (KANA, Kyo); Nakashinkawa-gun, Tateyama- cho, Mt. Tateyama, S. Matsuda s.n., Aug. 1893 (kyo), G. Nakai 4000 (kyo): Higashitonami- gun, Kamitaira-mura, Bunao Pass, 7. Mimoro & S. Tsugaru 11846 (Mo, TNs), G. Murata 30018 (KYO, TNs); Higashitonami-gun, Kamitaira-mura, Hosoo Pass, N. Mino s.n., 21 May 1964 (KANA); Shimoshinkawa-gun, Unazuki-cho, Mt. Karamatsu, Baba-dani valley, G. Murata & T. Shimizu 1904 (Kyo), J. Ohwi 7107 (Kyo), S. Okamoto s.n., 23 July 1935 (Kyo); Higashitonami-gun, Nawaga-ike Pond, N. Satomi 10150 (KANA, MAK, TNS). al. 22281 (GA, Gu); Yamagata city, Yamadera, K. Doi s.n., 10 Aug. 1966 (TNs), H. Ohashi s.n., 20 May 1960 (rus); Higashitagawa-gun, Tachikawa-machi, Mt. Gassan, R. Endo s.n., 28 July 1914 (rus); Nishimurayama-gun, Nishikawa-machi, Mt. Asahi-dake, Hara sn, 7 July 1959 (11); Nishiokitama-gun, Oguni-machi, Mt. lide, M. Ito 407 (Kyo, Mo), G. Koidzumi s.n., 1S Aug. 1910 (11), H. Ohba et al. 73071 (a, T1. rus), T. Yamazaki sn, 3 Aug. 1943 (11), 7. Yamazaki et al. 15 (11); Higashitagawa-gun, Asahi-mura, Kamina, K. Mort s.n., 30 July 1950 (11); Nishiokitama-gun, Oguni-machi, Oishisawa, 1984] YING ET AL., DIPHYLLEIA 89 S. Terabayashi 828 (kyo), Akumi-gun, Yuza-machi, Mt. Chokai, Univ. Tokyo Staff’s.n., 28 July 1888 (11); Nishimurayama-gun, Nishikawa-machi, Mt. Yudono, Univ. Tokyo Staff s.n., 23 July 1882 (G, T1); Yonezawa city, Mt. Azuma-san, Yoshida s.n., 6 Aug. 1915 (mak). Yamanashi Prefecture: Kitakuma-gun, Shirasu-cho, Mt. Kanno-dake, S. Kitamura s.n., 28 July 1931 (kyo), H. Matsuda s.n., 22 July 1954 (11), Y. Ogura s.n., 28 July 1920 (11); Kitakuma-gun, Shirasu-cho, Mt. oe take, G. Murata 11976 (KYO); Nakakuma-gun, Ashiyasu-mura, Mt. Kita-dake, H. Terao 895 (Kyo); “Fudzi yama” [Mt. Fuji-san], Tschonoski s.n., 1864 (Bm, p, Ups, W). Kuril Islands. KUNASHIRI: without collector or number, 1935? (kyo). U.S.S.R. SAKHALIN. Without further locality: Augustinowicz 18591 (BM), Augustinowicz s.n. (G, GH, Kk). Maoka Province: Maoka-gun, Maoka-san, 7. Mivake s.n., 4 June 1907 (sap); Katada-gun, Mt. Nota-san, 7. Mivake s.n., 28 June 1907 (sap). Motodamari Province: Motodamari-gun, Mt. Tasso-san, /. Nagai & S. Iwadera s.n., 23 July 1927 (sap). Odomari Province: Nagahama-gun, Shire- toko-mura, Mt. Shiretoko, 7. Miyake s.n., 11 July 1908 (sap, TNs). Shikika Province: Sanko-gun, Funadomari, !. Tatewaki 22705, 22961 (Kyo, sap). Tomari Province: Na- yoshi-gun, Nayoshi-mura, Kitakosawa, /. Henmi 63 (MAK). Toyohara Province: To- yohara-gun, Mt. Suzuya-dake, 7. Miyake s.n., 27 July 1907 (sap), M. Sato s.n., 19 July 1932 (71). We have seen no collections of Diphylleia from the Amur region of the Soviet Far East. The reports of its occurrence there by Kumazawa (1930), Li (1947), and others appear to be based on the title of the publication in which D. grayi was first described (Reisen in Amur-lande und auf der Insel Sachalin), rather than on actual specimens. Fedchenko (1937), in his treatment of the Berber- idaceae for Flora SSSR, attributed Diphylleia to only Sakhalin in the Soviet Union. Takeda (in Takeda & Tanabe, 1931) named plants with very deeply lobed leaves Diphylleia grayi var. incisa, and those with the lobes essentially lacking var. rotundata. The degree of lobing of the leaves in D. grayi is variable and continuous; plants with both deeply lobed and nearly unlobed leaves can be found intermixed in single populations. Kanai 742732 represents plants with lobed and unlobed leaves. There seems to be no basis for the recognition of these plants as infraspecific taxa. Takeda (in Takeda & Tanabe, 1931) did not cite specimens or indicate types in his descriptions of Diphylleia grayi vars. incisa and rotundata, and we have seen no specimens annotated or collected by him with these varietal names. In lieu of specimens, we wish to designate the illustrations accompanying the original descriptions of these two varieties as the lectotypes. In the more than 200 specimens of Diphylleia grayvi that we examined, seven had upper petioles longer than 1 cm (1.2 cm, 1.3 cm, 1.4 cm, 1.6 cm, 2.1 cm, 4.4 cm, and 5.8 cm). The most logical explanation for these aberrant plants is that although they have only two leaves, they represent forms transitional between plants with two and plants with three leaves. In the occasional plants with three leaves, the uppermost leaf is sessile or subsessile while the next lower leaf is petiolate. ACKNOWLEDGMENTS We wish to thank R. Mochizuki for the use of specimens in his personal herbarium, and the curators and directors of the following herbaria who made their specimens available for our study: 4, Akita Prefecture Museum, BM, DUKE, 90 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 E. EHUV, FARM, FI, G, GA, GH, GSMNP (Great Smoky Mountains National Park), HIB. K, KANA, KUN, KYO, L, LE, MAK, MO, NAS, NCSC, NCU, NwTc (Northwestern Teacher’s College, Wugong, Shaanxi, People’s Republic of China), NY, P, PE, SAP. TENN, Tl, TNS, TUS, TUSG, UPS, USCH, VDB, VPI, W, WCUH, WH, WU, WUK. We are grateful to S. Y. Hu, J. LaFrankie, S. A. Spongberg, and P. F. Stevens for carefully reading and commenting on the manuscript, and to A. M. Harvill, Jr.. and P. S. White for providing information on the distribution of Diphylleia cymosa. We also express our thanks to C. Z. Ji for preparing the illustrations of Diphylleia, and to K. Ueda and S. Y. Oh for supplying a photocopy of the paper by Lee. This material is based upon work supported in part by the National Science Foundation under Grant DEB-8119209 to P. H. Raven. This support, along with funding from the Arnold Arboretum of Harvard University and the Missouri Botanical Garden, allowed T. S. Ying to spend one year in the United States. LITERATURE CITED Airy SHaw, H. K. 1965. Diagnoses of new Paria new names, etc. for the seventh edition of Willis’s Dictionary. Kew Bull. 18: 249-273. Anonymous. 1974. Diphylleia. (In Chinese.) FI. a ee 1(2): 330, 331. Boarp oF EpucaTION OF Iwate PREFECTURE. 1970. Flora of Iwate Prefecture. (In Japanese.) Board of Education of Iwate Prefecture Bureau, E. 1904. 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A. 1980. eee of the Berberidaceae with an evaluation of classi- fications. Syst. Bot. 5: 149-17 92 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Micuaux, A. 1803. Flora Boreali-Americana. 2 vols. Levrault, Paris & Strasbourg. Miyaci, Y. 1930. Beitrage zur Chromosomenphylogenie der Berberidaceen. Planta 11: 650-659. MuRAKAMI, T.. & A. MATSUSHIMA. 1961. Studies on the constituents of Japanese Podophyllaceae plants. (In Japanese.) J. Pharm. Soc. Japan 81: 1596-1600. Muramatsu, H. 1932. Flora of Akita Prefecture. (In Japanese.) The Normal Schoo of Akita Prefecture Press. Nopa, M. 1969. Flora of Echigo Province in mid-Japan facing the Japan Sea. (In Japanese.) Vol. 2. Niigata University Press. Nopa, S., & T. Fusimura. 1970. Karyotypes in root-tip cells and endosperm nucleus of Diphylleia grayi. (In Japanese, English summary.) Kromosomo 79, 80: 2548- 2551. — Nocucut, J., & 8S. KAwWANOo, 1974. Brief notes on the chromosomes of Japanese plants. J. Jap. - 76-86. Nowicke, J. & J. J. Skvar_A. 1981. Pollen morphology and phylogenetic rela- tionships woe Berberidaceae. Smithsonian Contr. Bot. 50: 1-83. Numata, M., ed. 1974. The flora and vegetation of Japan. Elsevier Publishing Co. Tokyo. . ‘AWAKE, & D. Irow. 1972. Natural and seminatural vegetation in Japan. Blumea 20: 435-514 OGANozova, G. G. 1974, Anatomical structure of leaf in Berberidaceae s. |. related to the taxonomy of the family. (In Russian.) Bot. Zhurn. (Moscow & Leningrad) 59: 1780-1794. Renper, A. 1913. Rosaceae. nm: C. S. SARGENT, ed., Pl. Wilson, 1: 47-7 Roze, E. 1891. Séance du 13 novembre 1891. Bull. Soc. Bot. France 38: 324, 325. SARGENT, C. S. 1916. Numerical lists (of woody plants). Pl. Wilson. 3: 463-578. Scumipt, F. 1868. Reisen im Amur-lande und auf der Insel Sachalin. Mém. Acad. Imp. Sci. Saint Pétersbourg 12(2): 1-224. ScHuctz, C. H. 1832. System Pflanzenreichs. A. Hirschwald, Berlin Soepa, T. 1942. On the chromosomes of Diphylleia grayi Fr. Schm. (In Japanese.) Jap. J. Genet. 18: 47-48. Spurr, A. R. 1969, A low- eae resin embedding medium for electron microscopy. J. Ultrastr. Res. 26: 31- Srarteu, F. A. & R. S. a AN. 1976. Taxonomic literature. Bohn, Scheltema & Holkema, Utrecht. SUGAWARA, S. 1937. Plants of oe (In Japanese.) Group for the Support of Botanical Research in Saghalien. Takepa, H., & K. TANABE. (931. Kozan shokobutsu shashin-jushu. Ajushashobo, Tokyo. TAKHTAJAN, A. L. 1969. Flowering plants: origin and dispersal. (C. Jerrrey, translator.) Oliver & ra Edinburgh. LIKIAN, 1972. Comparative anatomical study of the seed coat anatomy of Leantice Gymnospermium, oe and allied genera in relation to their systematics. (In Russian.) Bot. Zhurn. (Mos w & Leningrad) 57: 1271-1278. oe ASHI, S. 1983. Studies in the mo een and systematics of Berberidaceae. . Floral anatomy of Diphylleia Michx. ae L. and Dysosma Woodson. ee Phytotax. Geobot. 34: 27-47, THiseELtoN-Dyer, T. F. 1889. CIX—vegetable productions, central China. Kew Bull. 1889; 225-227. Tuorne, R. 1968. Synopsis of a putatively phylogenetic classification of the flowering plants. Aliso 6: 57-66. TiscHLER, G. 1902. Die Berberidaceen und Podophyllaceen. Bot. Jahrb. Syst. 31: 596- 727. 1984] YING ET AL., DIPHYLLEIA ee ToOYOKUNI, H., & Y. ToyoKuNI (KIMURA). 1964. Ein neuer Anhalt fiir die Teilung der Podophyllaceen in zwei Unterfamilien. Bot. Mag. Tokyo 77: 197, 198. Woop, C. E., Jk. 1972. Morphology and phytogeography: classical approach to the study of disjunctions. Ann. Missouri Bot. Gard. 59: -124. Worsbe_Lt, W. C. 1908. A study of the vascular system in ee orders of Ranales. Ann. Bot. (London) 22: 651-682. ZHONG Guo YIXUIGEXUI YUAN Yowu Liansuoso. 1979. Zhong Yaozi. Vol. 1. People’s Health Press, Being. APPENDIX 1. Voucher specimens of Diphylleia used in anatomical studies. TAXON VOUCHER D. cymosa . 5. A., North Carolina, Avery County, FE. W. Wood & Bouf- fa 4150a (BM, CM, KYO , North ae Watauga County, Boufford & E. | oF 20954 (CM, KYO) D. grayi Japan, Se Pte Prefecture, Mt. Hyonosen, 8. Teraba- vashi 60 (Ky Ja apan, Honshu, Ishikawa Prefecture, Mt. Haku-san, S. Teraba- yashi Se 74 (KYO) apan, Honshu, as Prefecture, Mt. Shirouma, S. Teraba- “rash 793 (KY pan, - nshu, Yaa Prefecture, Mt. lide, M. /to 40 (Kyo) ae Hubei Province, Shennongjia Forest District, Wan Ji- D. sinensis ae Gale in ik no voucher collected) APPENDIX 2. Voucher specimens of Diphylleia used for pollen observations. TAXON VOUCHER U.S. A., North Carolina, Watauga County, Potato Hill, N of jet. of state roads 1324 and 1306, Kologiski & Perino 165 (Gu) U.S. A., Virginia, Smyth County, NE slope of White Top Mt. J. K. Small s.n., 28-29 May 1892 (GH) U.S. A., North Carolina, Macon County, 13 mi N of U.S. 64 on Rainbow Springs—Aquone Forestry Rd., Radford 5239 cu) D. cymosa D. grayi Japan, Honshu, Iwate Prefecture, Mt. Hakkoda, 4. Kimura & S. Sugaya s.n., 3 July 1952 (tus) Japan, es Miyagi Prefecture, Mt. Izumi, K. Saito & T. Kaneko s 18 May 1961 (Tus) Japan, ont Nagano Prefecture, Minamiazumi-gun, Hf. Ta- er Ai 366 (Tus) D. sinensis China, Gansu Province, “SE Gansu,”’ Licent 5325 (Bn China, Sichuan Province, Mealo Xian, Lysugou, Nan- ne Bei- mee Team 7164 PE) China, Yunnan Province, “Ma-cul-chan,” Delavay 3862 (A) —_ 94 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 T.S. Y. S. T. INSTITUTE OF BOTANY DEPARTMENT OF BOTANY ACADEMIA SINICA FACULTY OF SCIENCI 141 Hst CHtn MEN Wat TA CHIE Kyoro UNIVERSITY BEUING, PEOPLE’S REPUBLIC OF CHINA Kyoro 606, — DEB Present addre TSUMURA ae ATORY 9-9, 1-CHOME, IZUMI-HONMACHI Komae City, Tokyo 201, JAPAN HARVARD UNIVERSITY HERBARIA 2? Divinity AVENUE CAMBRIDGE, MASSACHUSETTS 02138, U.S.A ADDENDUM. Lectotypification of Diphylleia grayi F. Schmidt. Since this paper went to press, one of us (D. E. B.) has had the opportunity to examine the following specimens from Sakhalin, U.S. S. R., that were cited by Schmidt (1868) e his original description of Diphylleia grayt: no further locality, Glehn s.n., beginnin of August 1860; Dui, F. Schmidt s.n., June 1860 (Schmidt did not indicate in the Se cae that his collections were among those used in the original description, but the specimen cited as “Dui Anf. Juni 1860” is probably this one), Glehn s.n., 21 July 1860; Arkai, Glehn s.n., 27 May 1861; Kussunai, Bry/kin s.n., May 1860, Brylkin s.n., 25 April 1861 (all at Le). There are two plants on the sheet collected by eae on 21 July 1860, and both are atypical of the vast majority of Diphylleia grayi in having a petiolate upper leaf. It is almost certain, however, that Schmidt based his pet ee of the fruits of D. grayi on this specimen, since none of the other specimens contains mature fruits. Of the other specimens, Bry/kin s.n., 25 April 1861, is a sterile plant, and the Schmidt specimen has no flowers and only very young fruits. Glehn s.n., beginning of August 1860, is a mixed collection oo an upper leaf and infructescence (berries lacking) from one plant and the upper portion of another plant with two leaves and flowers, which must have been eda in the spring and not on the date indicated on the label. The specimen collected at Kussunai, Bry/kin s.n., May 1860, has only a single late flower and several very immature fruits and could not have been the source for Schmidt’s description of floral characters. The remaining sheet, Glen s.n., 27 May 1860, collected at Arkai, contains two very young plants with flowers in various stages of development; these were clearly the basis for Schmidt’s comments on the sepals, and for his comparison of them with those of D. cymosa Michaux, in his original description. There is also a small packet on this sheet that contains what appear to be portions of the roots of one of the plants. Of the two specimens on this sheet, the one on the right is the more complete, and it is this one that I wish to designate as the lectotype of a gray F. Schmidt. Photographs of the lectotype have been deposited at 4, Kyo, and I am grateful to the director and curators at Le for making these specimens available to me on loan.—D. E. B 1984] PAGE, LOISHOGLIA BETTENCOURTII 9 A POSSIBLE MAGNOLIOID FLORAL AXIS, LOISHOGLIA BETTENCOURTH, FROM THE UPPER CRETACEOUS OF CENTRAL CALIFORNIA VIRGINIA M. PAGE THE FOSSIL SPECIMEN described below was collected from an outcrop of Upper Cretaceous marine sediments in the foothills of the Diablo Range on the western border of the San Joaquin Valley near the central California town of Patterson. This locality, locally known as Black Gulch, has yielded numerous specimens of petrified woods (desenped in Page, 1967, 1968, 1970, 1973, 1979, 1980, 1981), as well as p ell-preserved pollen, spores, dinoflagellates, foraminifera, and radiolarians. eee to Bishop (1970), the sampled sequence at Black Gulch is correlated with zone D-2 of the forami- niferal zones established by Goudkoff (1945) and is equivalent to the Ragged Valley Shale member of the Moreno Formation. Due to a discrepancy in opinion regarding the position of the Maastrichtian-Campanian boundary with respect to the foraminiferal zones of Goudkoff, there is a lack of agreement as to the age of the D-2 zone, and hence of the Black Gulch locality. Whereas most workers place the boundary between Goudkoffs D-2 and E zones, Bishop (7m Chmura, 1973) stated that he believes the boundary to occur very near the D-1 and D-2 division. Since the Black Gulch locality lies within the D-2 zone, it is Maastrichtian 1n age according to most workers, but Campanian according to Bishop. Chmura (1973), who analyzed angiosperm pollen obtained from concretions, accepted Bishop’s judgment, but Stein (1983) found that the con- cretions contain a typical Maastrichtian assemblage of dinoflagellates. The opinion of the majority of workers is adopted here. The specimen is a calcareous petrifaction that measured approximately 3.5 cm in diameter and 3 cm in length before it was sectioned. Most of the tissues had been extensively altered, partly by pressure but chiefly by microorganisms, prior to fossilization. The general morphology is that of a herbaceous stem: a pith surrounded by a ring of vascular bundles and a cortex (FIGURE 1). The epidermis was not preserved. The pith is about 1.5 cm in diameter. Although much of the center has been destroyed, the cells—except for a peripheral zone of mostly small, relatively thick-walled ones—appear to have been large and thin walled. Numerous large cells averaging 102 um in diameter occur indi- vidually in the peripheral zone; each is surrounded by small, flattened cells. Lenticular plates of sclereids are abundantly scattered throughout. As seen in longitudinal section, these plates are about | cm long and 0.5 cm thick. It is difficult to determine the shape of individual sclereids, for they are closely packed and variously convoluted (FiGURE 2). Some, at least, are branched, esident and Fellows of Harvard College. 1984. Journal of the Arnold Arboretum 65: 95-104. ee 1984. 96 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Figure 1. Loishoglia bettencourtii: diagram of quadrant in transverse section show- ing arrangement of vascular bundles, x 6. Those aligned at periphery of pith are stelar. Note enlarged fascicles in various stages of division and departure from stele. Stippled portions of large fascicles denote phloem fibers and crushed phloem cells. Circles of bundles in cortex show variety of patterns. Disrupted bundle circles and atrophied bun- dles appear as outlines, nests of sclereids as outlines with stippled borders. Pith sclereids ath. core ‘ not shown. P = phloem cavities, S = “sheath while others have terminal bulges. In some cases, a mass of sclereids, as seen in transverse section, lies in close proximity to a protoxylem point ofa vascular bundle and extends into the interfascicular region. Individual sclereids within these masses are often elongated and lie parallel to one another (FiGuRE 3). They may be as much as 300 um in length. Although there is evidence ofa cambium within the vascular bundles, there is no discernible sign of an in- terfascicular cambium. Metaxylem vessels are by far the most conspicuous components of these bundles. The cells are comparatively large in diameter, averaging 45 um (range, 36-51) radially and 41 wm (28-50) tangentially, but they are less than 300 um in length. As far as can be observed, thickening of the walls of the conducting cells of the primary xylem is exclusively spiral. It is extremely difficult to interpret the nature of the thickening in the endwalls of the conducting elements in the area of overlap between elements. David Bierhorst examined the thin sections and was able to detect areas in the endwalls 1984] PAGE, LOISHOGLIA BETTENCOURTII OF of some of the elements where the spirals were diminished in thickness and resembled bars of scalariform perforations. The nonvascular tissue of the pri- mary xylem is composed of small cells, most of which have thick walls. Some of these cells may be parenchyma, while others appear to be fibers. Vessels in the secondary xylem are angular, thin walled, and mostly solitary, averaging 53 um (range, 36-57) in radial and 38 um (21-54) in tangential diameter. The length of the vessel elements is almost impossible to measure because cell outlines are very faint; also, since the vascular bundles are seldom oriented parallel to the plane of section, few well-defined and undistorted elements can be observed. The vessel elements are, therefore, merely estimated to be about 260-360 um long. Intervessel pits, in the few places where they can be observed, are small and opposite. Perforation plates are scalariform, with up to 20 but usually less than 10 bars. Parenchyma 1s sparse and occurs as single cells associated with vessels. Pits in the walls of parenchyma cells are large and lenticular. Medullary rays are heterocellular and 2 or 3 cells wide. Rays within vascular bundles are uniseriate. Thick-walled fibers with slitlike apertures make up the remainder of the nonvascular tissue. A large cavity surrounded by crushed cells occupies the presumed position of the phloem. Each bundle is capped by a group of narrow fibers. Within the ring of numerous stelar bundles in a transverse section, as many as 15 are much enlarged. Various stages of lateral expansion of the bundles— apparently a result of radial division—can be observed. The larger of these complexes comprise several segments of primary xylem, secondary xylem, and phloem and are separated from their neighbors by bi- or triseriate rays. The enlarged bundles are displaced outwardly into the cortex, and the position they occupied in the stele is filled with parenchyma. The ground tissue of the cortex is composed of large, relatively thick-walled parenchyma cells. Nests of scler- eids and large, isolated cells like those in the periphery of the pith are scattered throughout. Most conspicuous and crowded within the cortex are tight circles of vascular bundles; these are as large as 2 mm in diameter and include up to 16 bundles (FiGure 4). Individual bundles vary in size. Most contain both primary tissue and small cells in radial files in the position of a cambium and/ or undifferentiated cambial products (FiGure 5). Radially aligned cells are absent in smaller bundles. Except in a few cases, large cavities remain in the presumed position of the phloem (FiGure 6). Each bundle is capped by a group of narrow fibers, and each circle of bundles appears to be bounded by a sheath of flattened cells. Possibly these latter cells were flattened by pressure exerted by centrifugal expansion of the vascular bundles. In some instances the circular arrangement is disrupted, particularly toward the outer edge of the sections. Examination of a series of four transverse sections shows that some of the bundles of a partially disrupted circle remain associated, but that others are twisting away. Fragments of bundles traverse the outer edge of the sections both vertically and horizontally. The arrangement of vascular traces in the cortex appears to be unique, for a search through the literature failed to reveal evidence of similar circles of bundles in the cortex of any structures of either monocotyledons or dicotyledons. I must point out, however, that complete anatomical studies of axes of the primary body of woody plants are very limited; — pot 98 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Ficures 2-6. Loishoglia bettencourtii: 2, nests of sclereids, transverse section, show ing convoluted pattern, * 100; 3, pith sclereids, longitudinal section, showing cieasated form, x 100: 4, circle of cortical bundles, transverse section, x 37; 5, enlargement of portion of 3 contiguous cortical bundles, transverse section, showing undifferentiated secondary xylem above and primary xylem with thick-walled vessels below, x 100 1984] PAGE, LOISHOGLIA BETTENCOURTII 32 studies of inflorescence axes of both woody and herbaceous plants are even more limited. SYSTEMATIC DESCRIPTION Loishoglia Page, gen. nov. Axes with central cylinder of collateral bundles surrounding a pith composed of parenchyma, nests of sclereids, and isolated secretory cells, and a cortex with narrowly wedge-shaped vascular bundles arranged in numerous tight circles of up to 16 bundles per circle. Metaxylem vessels with walls spirally thickened, perforations scalariform. Vessels of secondary xylem solitary with small, op- posite intervessel pits. Perforations scalariform with up to 20 but usually fewer than 10 bars. Medullary rays heterocellular, 2 or 3 cells wide. Type species. Loishoglia bettencourtil Page. Loishoglia bettencourtii Page, sp. nov. Pith. 1.5 cm in diameter. Parenchyma cells large, thin walled except for pe- ripheral zone of small, thick-walled cells. Secretory cells avg. 102 um in di- ameter. Sclereids in lenticular plates | cm long and 0.5 cm thick in longitudinal section. Vascular bundles. Vascular bundles numerous. Primary xylem vessels solitary, isodiametric, avg. 45 wm (range, 32-50) in diameter, ca. 300 um long. Perfo- rations scalariform. Secondary xylem vessels mostly solitary, avg. 53 um (36- 57) in radial diameter, 38 wm (21-54) in tangential diameter. Vessel element length ca. 360 um. Intervessel pits small, opposite. Perforations scalariform, with up to 20 but usually fewer than 10 bars. Parenchyma sparse, occurring as single cells associated with vessels. Medullary rays 2 or 3 cells wide, het- erocellular; rays within bundles uniseriate. Ground tissue of thick-walled fibers with slitlike apertures. Phloem fibers thick-walled, narrow, forming centrifugal cap as viewed in transverse section, sieve cells not preserved. Cortex. Circles of vascular bundles numerous; individual bundles similar to those of central stele except secondary tissues undeveloped or lacking. Solitary bundles numerous at periphery. Ground tissue parenchymatous with numerous nests of sclereids. Secretory cells abundant. MATERIAL. One calcified petrifaction measuring 3.5 cm in diameter and 3 cm in length. Ho ortype. California Academy of Sciences Geological Collections no. 61208. individual vascular bundle, transverse section, showing primary xylem to right and cap of phloem fibers to left of large phloem cavity (note crushed cells surrounding cavity), x 70. 100 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 The genus is named after Lois Hogle in recognition of her kindness in providing transportation to the collection site. The specific epithet 1s in honor of the Bettencourt family of Crow’s Landing, California, for allowing me to collect on their property over the years. Loca.ity. Moreno formation. Black Gulch; NE%s, SE sect. 32, R7E, T5S, Oristimba 15” quadrangle, California. AGe. Maastrichtian. DISCUSSION The specimen remained an enigma for a long time mainly because of the unusual structure of the cortex and the difficulty in determining the nature of the endwalls of the vessel elements in both the primary and the secondary xylem. The presence ofa fascicular cambium is an indication that the specimen is a dicotyledon, for in monocotyledons the cambium 1s exclusively extrafas- cicular. The anatomy of the stele has characteristics of certain dicot stems. The numerous gaps left by traces in various stages of departure from the stele are suggestive of a system of much-telescoped internodes and possibly a spiral phyllotaxy such as one would expect to find either in the receptacle of a lower with numerous parts or in the peduncle of an inflorescence. Among flowers examined, receptacles of members of the Magnoliaceae (par- ticularly in the perianth region in Magnolia L. and Liriodendron L.) proved anatomically most similar to the fossil. In plants of both Magnolia and Lir- seen on, numerous collateral bundles surround a large pith containing nests of sclereids and secretory cells. Cambial activity is minimal and confined to the oo bundles. Sclereids and secretory cells occur in the stems of many genera of dicots, but the combined occurrence of nests of sclereids of the type observed in Loishoglia and secretory cells in both pith and cortex is not com- mon and, as far as | can determine, has been reported only in certain families of the Magnoliales. Among these families, vessel elements of the type observed in the secondary xylem of the fossil are found only in genera of the Magnoli- aceae. Although much larger than the secretory cells of the species of Magnolia, Michelia L., and Liriodendron examined, the enlarged cells of parenchymatous tissues in Loishoglia may be homologous with them, for they are similar in shape and distribution. In the modern forms examined these cells usually are not surrounded by flattened parenchyma cells as they are in the fossil. It is possible that the condition in the fossil 1s due to excessive expansion of the secretory cells early in differentiation. Although of the same general type as those in the Magnoliaceae, sclereids in Loishoglia tend to be much longer. The structure of the vascular elements in the fossil is consistent with that in many members of the Magnoliaceae. Scalariform perforations with less than ten bars are characteristic of vessels in the secondary xylem, and opposite intervessel pits are present in a number of genera. Vessel elements of the primary xylem range from slightly over 200 «m to 500 um in length in the stalk and receptacle of the magnolia flowers examined. Because of poor preservation, 1t was 1m- possible to measure more than one vessel in the primary xylem of the fossil. 1984] PAGE, LOISHOGLIA BETTENCOURTII 101 This element, which was 290 um long, falls in the lower portion of the range observed in Magnolia. However, the numerous fragments visible in longitu- dinal sections indicated that the average length of vessel elements was probably greater than 290 um. The most obvious difference between the fossil and the modern forms lies in the cortex. Most of the cortical bundles in the receptacle of Magnolia or Liriodendron are single or in pairs, but occasionally a circle of three (or rarely four) occurs, particularly in Liriodendron. In the specimens examined (L. tulipifera L. and ornamental varieties of M. grandiflora L.) cor- tical bundles occurring in circles are broadly wedge shaped and arranged (as in the fossil) so that in transverse section the apices of the wedges are in juxtaposition. Parenchyma cells immediately surrounding a circle of bundles differ only slightly in shape and size, so there is nothing resembling the sheaths of flattened cells characteristic of the circles of bundles in the fossil. If the fossil is homologous with floral axes of members of the Magnoliaceae, an explanation of the pattern of cortical traces 1s necessary. Two interpretations are possible. First, the specimen may represent a portion of the axis of an individual flower, and the cortical bundles traces to its various components. As noted above, occasional circles of three or four wedge-shaped vascular bundles occur in transverse sections of flower stalks and receptacles of Magnolia and Liriodendron. It is probable that these circles represent situations in which branches of traces developed parallel to one another for a short distance before diverging from the parent trace. Although more complex, the condition in the fossil may be similarly explained. The seeming complexity may stem from the more profuse branching of traces in the fossil. In sections of receptacles of Liriodendron and Magnolia, many traces diverging from the stele appear to be dividing bilaterally. In the fossil some stelar bundles appear to be dividing bilaterally, but others look as though they were undergoing multiple divisions (see Ficure 1). Whereas in the modern forms these branches soon diverge, in the fossil most remain associated for at least the length of the specimen, and some are joined by a sympodium from another level of the stele. (Since the direction of provascular development is not known, trace patterns are discussed here as if they had been basipetal.) The second interpretation, derived from Nast’s (1944) description of the vasculature of the cymelike inflorescences in species of Drimys J. R. & G. Forster (Winteraceae), is that the specimen may have been part of the bedancle of a compound inflorescence, in which case the circles of bundles may be traces to pedicels. Nast (p. 458) stated that in the flowering stalk, “each thickened area of the stele breaks up into a number of bundles as it leaves the central cylinder and almost immediately assembles into steles of the floral pedicels. The pedicels may contain one or two concentric bundles or a cylinder of bundles.” The cortical circles of bundles in the fossil, Loishoglia, may be homologous with the pedicel traces of Driniys; however, unlike the latter, which enter the pedicels at more or less the same level in the axis as they depart from the central stele, those in the fossil continued on at a much more acute angle for some distance before connecting with a pedicel. That groups of traces may have maintained a circular configuration on departure from the main axis 1s suggested by the presence of circles in longitudinal sections tangential to the LOZ JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 outermost edge of the specimen. More difficult to explain in either interpre- tation are the numerous individual bundles and small circles crowded at the outer edge of the cortex in transverse sections. In these bundles the cells are crushed and wrinkled as if atrophied. Although Loishoglia has several anatomical characteristics in common with the Magnoliaceae, the unique vascular pattern of the cortex distinguishes it from all members of both that family and related ones. If the circles of bundles in the cortex represent incipient steles of pedicels, this would suggest that the inflorescence was complex. Although flowers of the Magnoliaceae are generally solitary, in E/merrillia Dandy and Michelia there may be more than one per inflorescence. Moreover, there is some evidence that the flowering axes of the closest relatives of the Magnoliaceae, the Degeneriaceae and Himantandraceae, may be reduced from more complex structures. In Ga/bulimima F. M. Bailey (Himantandraceae) bracts at the base of the receptacle subtend axillary buds that occasionally develop into subsidiary flowering axes (Bailey ef a/., 1943). The flowering axes of Degeneria I. W. Bailey & A. C. Sm. bear bracteoles thought by Bailey and Smith (1942) to indicate that the axes had been reduced from more complicated systems. Compound inflorescences, therefore, may have been characteristic of the ancestral group from which these families were derived. Three wood and stem specimens showing possible affinities with the Mag- noliaceae or related families have previously been described from the Maas- trichtian of central California. Two of these (suptc! 10,000, Magnoliaceoxylon 5 (Page) Wheeler, Scott, & Barghoorn (Page, 1970), and casG 60425 (Page, 1980)) are eer of secondary wood, and one (cAsG 60133 (Page, 1980)) 1s a young stem with pith and both primary and secondary xylem. The possible relationship between the latter specimen and the Magnoliaceae was not recognized initially because its solitary pores and heterocellular rays are not typical of the family. It does, however, contain pith sclereids of the mag- noliaceous type, and secretory cells are present in the rays and pith. Of the three specimens, 60133 is most similar anatomically to Loishoglia. It 1s a stem or branch 2.5 cm in diameter with a large pith | cm in diameter. Pores in the metaxylem are 21-43 um in diameter; those in the secondary xylem 86-96 um. Intervessel pits in the secondary xylem are small and opposite, and per- foration plates are scalariform with less than ten bars. Secretory cells in the pith are devoid of contents and difficult to distinguish in transverse section from the very large, thin-walled parenchyma cells, but a few can be observed in the longitudinal plane where the parenchyma cells are mostly flattened. The secretory cells are comparable in size to those in Magnolia (avg. 80 wm) and are thus smaller than those observed in Loishoglia. Enlarged cells in the rays, interpreted as secretory (Page, 1980, fig. 4), are closely similar to such cells in the rays of secondary xylem in Magnolia. Pith sclereids are much like those in Loishoglia. An additional feature typical of the Magnoliaceae can be ob- served in transverse sections of specimen 60133. The sections are through a ‘Stanford University Paleontological Type Collection, now housed at the California Academy of Sciences, San Francisco 1984] PAGE, LOISHOGLIA BETTENCOURTII 103 multilacunar node with a large median trace flanked by two small traces and eight additional lateral traces. As described by Canright (1955), the typical nodal pattern in Magnolia is similar except for the presence of a stipular trace opposite the median trace. Tissues in the area where a stipular trace would be expected were not preserved in the fossil; therefore, it is not possible to deter- mine whether such a trace was originally present. Several traces are present in the fossil that are not figured in Canright’s diagram. This difference may not be significant, for he has pointed out that the number of nodal traces in species of Magnolia is extremely variable both inter- and intraspecifically; however, although the number of lateral traces may vary from four to fifteen, the median and stipular traces are always present. The numerous anatomical similarities observed in cAsG 60133, Loishoglia, and stems of the Magnoliaceae suggest that the fossils were related to one another and to the Magnoliaceae. Although vessel elements in the secondary xylem of Loishoglia are on the average smaller in diameter than those in either 60133 or most members of the Magnoliaceae, they compare well in size with elements in the earliest-formed secondary xylem (where the vessels are about half the diameter of those in subsequent increments). Secretory cells in Lovs- hoglia, however, are much larger than those in 60133 and the species of Mag- nolia, Michelia, and Liriodendron examined. It is possible that the two fossils were related but not necessarily derived from members of the same natural species or genus. The locality from which 60133 was collected is slightly younger than the horizon in which Loishoglia was found. It has been estimated (Page, 1981) that the two localities are separated by an interval of about 1.5 million years. The two specimens of secondary wood, suptc 10,000 (from the same locality as Loishoglia) and casG 60425 (from the same locality as 60133), are closely similar to the woods of the modern Magnoliaceae except for the absence of terminal parenchyma. They differ from each other in that the intervessel pits appear to be scalariform in 60425, whereas they are mostly opposite in 10,000. Neither specimen shows evidence of secretory cells, and since both consist only of secondary wood, no sclereids are present. Until additional material becomes available, it is useless to speculate on the relationship between these two spec- imens and Loishoglia. The lack of secretory cells in itself may not be significant: among modern species of Magnoliaceae it is not unusual for such cells to be absent from the secondary wood but consistently present in primary tissues. The magnolialian complex can be traced at least to the early part of the Upper Cretaceous, as indicated by fruits and leaves described by Dilcher ef al. (1976), Crane and Dilcher (1982), and Dilcher and Crane (1982, 1984) from the early Cenomanian of Kansas. When compared with modern derivatives of the ancestral complex, Loishoglia appears to be most closely allied to the clade consisting of the Degeneriaceae, Magnoliaceae, and Himantandraceae. Lois- hoglia and the putative magnolioid wood specimen (cAsG 60133) can be added to a growing list of extinct Cretaceous genera of dicotyledons. The knowledge that a significant number of Cretaceous angiosperm genera are now extinct should be sufficient reason for caution in assessing the degree of relationship between pre-Tertiary woods and supposed modern counterparts. 104 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 LITERATURE CITED Baitey, lL. W., & A.C. Smit. 1942. Degeneriaceae, a new family of flowering plants from Fi. ee Arnold Arb. 23: 356-365. . Nast. 1943. The family Himantandraceae. Jour. Arnold Arb. 24: 190- aA Bisuop, C. 1970. Upper Cretaceous stratigraphy on the west side of the northern San Joaquin Valley, Spon and San Joaquin counties, California. Calif. Div. Mines Geol. Spec. Rep. 104. CANRIGHT, J. E. 1955. - comparative morphology and relationships of the Mag- noliaceae, [1V. Wood and nodal anatomy. Jour. Arnold Arb. 36: 119-140. Cumura, C. A. 1973. Upper Cretaceous (Campanian—Maastrichtian) angiosperm pol- len ee the western San Joaquin Valley, Calif., U.S.A. Palaeontographica 141B: 89-17 CRANE, P., : D. Di-cHer. 1982. Magnoliid fruiting axes ey ae mid-Cretaceous of North America, II. (Abstract.) Bot. Soc. Am. Misc. Publ. DL Ditcuer, D., & P. CRANE. 1982. 5 aa fruiting axes A - mid-Cretaceous of North America, I. (Abstract.) Bot. Soc. Am. Misc. Publ. 162: 5 & 1984. Archaeanthus: an early angiosperm from the Cenomanian of the western enor of North America. Ann. Missouri Bot. Gard. (in press). Crepet, & C. D. Beeker. 1976. Reproductive and vegetative morphology ofa Hone angiosperm. Science 191: 854-856 GoupbkorF, P. P. 1945. Stratigraphic relations of Upper eerie in Great Central Valley, California. Am. Assoc. Petrol. Geol. Bull. 29: 956-1007. Nast, C. G. 1944. The ee morphology of the wee VI. Vascular anatomy of the flowering shoot. Jour. Arnold Arb. 25: 454-465. Pace, V.M. 1967. Angiosperm ar from the Upper Cretaceous of central California: so Am. Jour. Bot. 54: 510-514. 8. a wood from the Upper Cretaceous of central California: part Il. bi. 55: 168-172. An uae ey wood from the Upper Cretaceous of central California: part HHI. tid 57: 1139- 973. A new ane from the Upper Cretaceous of central California. /bid. 60: il 575 1979, Dieotyledonous wood from the Upper Cretaceous of central California. Jour. Arnold Arb. 60: 323-349. 980. Dicotyledonous wood from the Upper Cretaceous of central California, If. oo 61: 723-748. 1981. oe ue from the Upper Cretaceous of central California, IIT. Conclusions. /bid. 62: -455 STEIN, J. 1983. oo ae mpanian—M ichtian) dinoflagell s fro the Great Valley Group, central Calfornis. 409 pp. Unpubl. Ph. D. ck Stanford University. DEPARTMENT OF BIOLOGICAL SCIENCES STANFORD UNIVERSITY STANFORD, CALIFORNIA 94305 1984] HAYDEN ET AL., PICRODENDRON 105 SYSTEMATICS AND PALYNOLOGY OF PICRODENDRON: FURTHER EVIDENCE FOR RELATIONSHIP WITH THE OLDFIELDIOIDEAE (EUPHORBIACEAE) W. J. HAYDEN, W. T. Gituist, D. E. STONE, C. R. BROOME, AND G. L. WEBSTER ALTHOUGH KNOWN to botanical science for 285 years, the genus Picrodendron Planchon has been poorly understood for most of this time. The most pervasive problem has been that of discerning familial relationships, and there have been additional difficulties in typifying the generic name (Hayden & Reveal, 1980) and in distinguishing its three nominate species. This paper provides a system- atic treatment for Picrodendron and demonstrates its relationships with Eu- phorbiaceae subfam. Oldfieldioideae Kéhler & Webster as evidenced by data on gross morphology, palynology, anatomy, and cytology. Picrodendron captured the attention of Dr. William T. Gillis while he was pursuing floristic studies of the Bahama Islands. During the decade preceding his death, Gillis accumulated a considerable amount of information on this problematic genus; he was attempting to complete this long-standing project in the weeks before he died in June, 1979. It has been a privilege tempered with sadness for one of us (W. J. H.) to prepare this paper from Gillis’s notes and the contributions that he had solicited from the other authors. Herbarium and field studies reported here were performed by Gillis and form the basis for much of what appears in the generic description and the sections on bio- geography and economic uses; taxonomic judgment at the species level is also Gillis’s. Broome and Stone contributed descriptions of pollen and most of the discussion concerning the Juglandaceae: information on relationships with the Euphorbiaceae was provided by Webster. This paper is a tribute and memorial to Gillis, an indefatigable student of the Bahama flora. MATERIALS AND METHODS Taxonomic treatment of Picrodendron is based on examination of numerous specimens in the wild, in cultivation, and in the herbarium. Herbarium spec- imens examined are listed in the ApPpENDIX. Pollen descriptions are based upon examination of the following specimens. Austrobuxus carunculatus (Baillon) Airy Shaw: New Caledonia, Quinne-Tal, Stauffer & Blanchon 5751 (puKe); New Caledonia, Kouébuni, Hiirlimann 571 (us). Austrobuxus eugeniifolius (Guillaumin) Airy Shaw: New Caledonia, Guil- President and Fellows of Harvard C ae 1984. Journal of the Arnold Arboretum 65: 105-127. January, 1984. 106 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 laumin & Baumann-Bodenheim 12908 (us). Mischodon zeylanicus Thwaites: Sri Lanka, De Silva 28 (us). Oldfieldia africana Bentham & Hooker f.: Ivory Coast, Voorhoeve 1963 (Mo). Picrodendron baccatum (L.) Krug & Urban: Cuba, Santiago, Clemente 6587 (Gu, us); Jamaica, Harris 12516 (us). Piranhea lon- gepedunculata Jablonski: Venezuela, Blanco 299 (us). Tetracoccus dioicus Par- ry: United States, California, Munz 12614 (mo). Tetracoccus hallti Brandegee: United States, California, Dressler 1211 (mo); Arizona, Kearney & Harrison 7530 (us). For brightfield microscopy, dried pollen was acetolyzed, treated with potas- sium hydroxide, and mounted in glycerine jelly. Since the pollen grains were nearly spherical, diameter measurements were made without regard to the polar axis: a minimum of 30 grains were measured at x 1000 with a x 40 high dry objective and = 25 oculars. Photomicrographs of acetolyzed grains were taken with a x 90 oil immersion apochromatic objective (n.a. 1.32) anda x 10 neg- ative ocular at x 1500 on Kodak Ortho or Pan sheet film, or at x 370 on Kodak 35 mm High Contrast Copy film. For SEM observations, acetolyzed pollen grains were air dried from 70% ethanol and then coated with gold- palladium (60:40) in a vacuum evaporator. For TEM, dried pollen was fixed for 10-12 hours in 2% glutaraldehyde buffered with 0.1 M Sorensen’s phosphate buffer at pH 7.0, washed, and finally postfixed for 2 hours in 2% phosphate- buffered osmium tetroxide. Immediately after stepwise dehydration in ethanol, the pollen was embedded in Spurr resin, standard formulation. Thin sections were poststained either in aqueous potassium permanganate solution or in a uranyl acetate-lead citrate combination. Observations of seedling morphology in Picrodendron were obtained from greenhouse-grown seedlings at the University of Richmond, Richmond, Vir- ginia. The seeds came from plants cultivated at the Fairchild Tropical Garden and originally collected from Driggs Hill, South Andros, Bahama Islands. Voucher specimens for these plants are Gillis 10924, 10975, and 11031] (AAH, FTG, IJ). TAXONOMIC HISTORY A century and a half elapsed between the first published record of Picroden- dron by Sloane (1696) as “Nux juglans trifolia. . .” and its ultimate recognition as a distinct genus (Planchon, 1846). It is not surprising, perhaps, that colonial Europeans, overwhelmed by the novelty and diversity of the neotropical flora, associated this plant with the familiar walnut, since there are certain gross resemblances. Consider, for example, the much-quoted but uncritical passage from Browne (1756) describing the “Jamaica walnut”: “The outward part of the fruit is soft and pulpy, when ripe; but the hard ligneous shell, and the partitions and lobes of the seed, as well as parts of the flower, agree perfectly with the general characteristics of the genus.”’ Linnaeus apparently saw neither living nor herbarium specimens of Picrodendron, which he nevertheless named Juglans baccata (Linnaeus, 1759), referring to Browne (1756), who had in turn referred to Sloane’s (1725) illustration (see FiGure 1). Amazingly, this link with the Juglandaceae has persisted for over two and a half centuries despite 1984] HAYDEN ET AL., PICRODENDRON \ N Bee ), jie i de ily Mla tie all ae H Lee / Nus priadaaas Peftvifodia, fructu Pdiviers moltehatne, | | Maciitidme Figure 1. Lectotype of Picrodendron baccatum (reproduced from H. Sloane, A Voy- age to the Islands Madera, Barbados, Nieves, S. Christophers and Jamaica 2: 1. 757, fig. 5-125 108 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 having been based on very superficial analyses and a decidedly nondefinitive illustration. Besides being considered a walnut, Picrodendron has also been treated as Tapia (i.e., Crateva L.) (Capparaceae), by Catesby (1734); Schmidelia (1.e., Allophylus L.) (Sapindaceae), by Richard (1845); and Rhus (Anacardiaceae) by Macfadyen (1837). Actually, Macfadyen had misapplied De Candolle’s (1825) name Rhus arborea (basionym: Toxicodendron arboreum Miller). The names of Miller and De Candolle apply to A//ophylus, but Macfadyen’s specimen unmistakably represents what 1s now known as Picrodendron. Macfadyen’s error unfortunately threw the application of Planchon’s (1846) name Picro- dendron into uncertainty, for Planchon based the genus on both an authentic Macfadyen specimen (k!) and the basionym Rhus arborea DC. Planchon’s species name, P. arboreum, is therefore a synonym of Allophylus L. Prior to the Sydney Congress of 1981, however, the Code of Botanical Nomenclature was unclear whether, in such a situation, the name Picrodendron should be typified by the species represented in the literature citation (1.e., Rhus arborea, Allophylus) or by the species represented by the specimen cited (1.e., Macfadyen s.n.). Consequently, Hayden and Reveal (1980) proposed to conserve Picro- dendron from the 1859 publication of Grisebach; full particulars can be found in that paper. However, revisions incorporated into the Code at Sydney (see Taxon 30: 103-105, 200-207, 904-911) now permit a different course. New Article 10.4 states, ““By conservation, the type of the name of a genus can be a specimen used by the author in the preparation of the protologue, other than the type of a name of an included species.” Further, new Article 14.8 reads, ‘“A name may be conserved with a different type from that designated by the author or determined by application of the Code.” Thus, we herein retain Planchon’s name, to be typified. by conservation, by Macfadyen’s specimen of Picrodendron baccatum. Picrodendron has been treated variously as a single species or as two or three insular endemics. Grisebach (1859) realized that the Jamaican Juglans baccata L. and Picrodendron arboreum Planchon were conspecific with the Cuban Schmidelia macrocarpa A. Rich. Grisebach called these plants Picrodendron juglans, a superfluous name that was also applied to plants from the Bahamas (Gardiner & Brace, 1889). Urban (1893) restored Linnaeus’s epithet, forming Picrodendron baccatum (L.) Krug & Urban for Jamaican and Cuban elements, but distinguished Bahamian material as var. bahamense Krug & Urban. Britton (1906) preferred to treat Cuban and Bahamian elements as one species, P. macrocarpum (A. Rich.) Britton. Small (1917) distinguished plants from Hispaniola as a third species, Picro- dendron medium Small, and his concepts of the three species can be summa- rized as follows: P. baccatum from Jamaica, characterized by dark leaflets with acute apices, long fruiting pedicels, and spherical fruits with thin endocarp; P. macrocarpum from Cuba and the Bahamas, characterized by blunt, pale leaflets, short fruiting pedicels, and somewhat ovoid fruits with thick endocarp; and P. medium from Hispaniola, characterized by leaflets similar to those of P. bac- catum, and endocarp similar to that of P. macrocarpum. Small’s distinctions seem trivial; moreover, these three epithets have been used frequently in senses 1984] HAYDEN ET AL., PICRODENDRON 109 other than those outlined by Small (Gémez de la Maza & Roig y Mesa, 1916; Urban, 1920; Barker & Dardeau, 1930; Moscoso, 1943; Howard, 1955; Adams, 1972). As noted above, early concepts of the relationships of Picrodendron varied widely, and disparate taxonomic associations have persisted throughout most of its subsequent history. Varying opinions regarding its relationships include placement in or association with the Simaroubaceae (Planchon, 1846; Bentham & Hooker, 1862; Urban, 1920: Barker & Dardeau, 1930; Moscoso, 1943), Juglandaceae (Grisebach, 1859), Burseraceae (Grisebach, 1866), Terebintha- ceae (Hallier, 1908), Picrodendraceae (Small, 1917, and numerous recent au- thors), Euphorbiaceae (Fawcett & Rendle, 1917: Thorne, 1976; Webster, 1975; Takhtajan, 1980; Cronquist, 1981), and Bombacaceae (Hallier, 1921). The family Picrodendraceae has been assigned to the Juglandales (Cronquist, 1968: Hutchinson, 1973), Rutales (Scholz, 1964; Takhtajan, 1966), and Euphorbiales (Novak, 1961; Takhtajan, 1969; Airy Shaw, 1973: Cronquist, 1978). ECONOMIC IMPORTANCE There are several recorded uses of Picrodendron, all of minor economic consequence. In Hispaniola the plants are commonly grown as a windbreak. In Jamaica the fruits are occasionally eaten, although Fawcett and Rendle (1920) stated that this should be done with caution. Leén and Alain (1951) reported that the leaves are eaten by animals. Curiously, one of its common names in the Dominican Republic, ““mata becerro,”” means “‘calf killer.”’ Ji- ménez (pers. comm.) reported that in the Dominican Republic leaves of Pic- rodendron are reduced to powder and used to kill bedbugs and lice. Sawyer (1955) reported that on Great Inagua a tea made from the leaves is used in the treatment of poisoning from eating fish; additional medicinal uses were reported in Roig y Mesa (1945). The viscid exocarps of Picrodendron fruits have been employed for their saponin in washing clothes. Record and Hess (1943) mentioned that the wood, which finishes smoothly and is resistant to decay, has been used in naval construction and turnery. SYSTEMATIC TREATMENT Picrodendron Planchon, London J. Bot. 5: 579. 1846, nom. cons. prop. Type: ‘“Jamaicae, Mac Fadyen, in herb. Hook..” typ. cons. [Juglans baccata L. =P. baccatum (L.) Krug & Urban]. Tree to 12 m, dioecious (rarely monoecious): bark rough, bitter to taste, gray, furrowed to shaggy in age: shoots finely pubescent. Leaves alternate, palmately trifoliolate, 1-2 dm long, deciduous prior to flowering: stipules inconspicuous, early deciduous; petioles 4-10 cm long, minutely pubescent; leaflets with the petiolules 0.5—2.5 cm long, joined at base, the blades 1.5-1 1 cm long, elliptic, oblong, ovate, or obovate, obtuse to emarginate apically, unequal basally on lateral leaflets, otherwise symmetrical, pinnately veined, finely reticulate, with the upper surface glabrous to glabrate and occasionally with tiny hairs proxi- mally on midvein, the lower surface slightly paler green than the upper. glabrate 110 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 to puberulent but densely pubescent on major veins. Flowering first during third year. Staminate flowers in stalked, puberulent catkins borne in axils of leaves of previous season’s growth, appearing before or with leaves in com- pressed heads, elongating to loosely clustered spikes; flowers greenish yellow, subtended by (1 to) 3 (to 7) bracts, these imbricate, to 1.5 mm long, with central one larger and overlapping laterals, acute, deltoid; perianth absent, stamens 3 to 54, on convex hemispherical receptacle, the filaments free, shorter than anthers (less than 0.5 mm), short-subulate, glabrous, the anthers 2-celled, ba- sifixed, to 1.5 mm long, oval, notched at apex and cordate at base, sparingly puberulent especially at tip, dehiscing longitudinally when mature, slightly extrorse: pollen spinose-verrucate, 5- to 8-porate; pistil absent. Pistillate flowers axillary, borne singly on shoots of current season; pedicels 2—4 cm long, abruptly widened below the finely pubescent, concave receptacle; flowers green, sub- tended by 4 or 5 valvate bracts, these ascending-recurvate, sepaloid, + unequal, 2.5-8 mm long, lanceolate, gland bearing, remotely toothed or ciliate; stam1- nodes absent; ovary superior, sessile, ovoid, 2-carpellate, naked; the style ter- minal, slightly longer than ovary, columnar, the stigmas 2, divergent, longer than style, stout-subulate, with revolute margins; ovules 2 per locule, pendulous from central axis, anatropous, the raphe ventral, the integuments 2, the ob- turator cushionlike, reddish brown, attached to placenta above ovules, covering micropyles, withering in fruit. Fruits smooth, 2-locular, indehiscent, globose to ovoid drupes, 1.5-2.5 cm long, yellow-orange to yellowish green at maturity, nodding, dry black; exocarp thin, fleshy, with numerous vesicles bearing bitter juice; endocarp 1.5-2 mm thick, woody to bony, brittle, marked by 4 equi- distant longitudinal lines. Seed(s) | (all but | ovule usually aborting) (rarely 2 with | in each locule), nearly terete to irregularly ovoid; endosperm lacking; testa membranaceous, enclosed between folds of cotyledons; hypocotyl and micropyle superior, epicotyl large, turned back on funicle at right angles to cotyledons; cotyledons plicate, corrugated; nucellus convoluted. A monotypic neotropical genus found on the Bahama Islands, Cayman Is- lands, Cuba, Hispaniola, Jamaica, and the Swan Islands (see Map 1). Picrodendron baccatum (L.) Krug & Urban, Bot. Jahrb. Syst. 15: 308. 1893. Juglans baccata L. Syst. Nat. ed. 10. 1272. 1759. Lectotype: Sloane, Voy. Isl. Madera, Barbados, Nieves, S. Christophers & Jamaica 2: 1. 157, fig. 1. 1725. The typotype is in the Sloane Herbarium, H.S. 5: 49 (pm!). Picrodendron juglans Griseb. Fl. Brit. W. Indian Is. 2: 177. 1859, nom. superfl. FIGURE 2. Schmidelia macrocarpa A. Rich. in Sagra, Hist. Phys. Pol. Nat. Cuba 10: 116. p/. 30. 1845. Type: Cuba, Sagra s.n. (lectotype, p, Herb. Rich.!). Picrodendron macrocar- pum (A. Rich.) Britton, Bull. New York Bot. Gard. 4: 139. 1906. I oe baccatum var. bahamense Krug & Urban, Bot. Jahrb. Syst. 15: 308. 1893. Type: Bahama Is., Hog Is. ue w Paradise Is., off New Providence], March [7], Eggers 4402 (holotype, B, destroy bee medium Small, Bull. New York Bot. Gard. 18: 185. 1917. Typr: Sant see a .Azua, northeast of Azua, 20 March 1913, Rose et al. 4042 (holoty i as pe, us!). 1984] HAYDEN ET AL., PICRODENDRON Lid on “\ a a) ee. Pa - = 7 si re) We _ ~ = S, act \ A ee Se ? oy \ a i eS < oe e e % « te: Su a | i i og _ ‘oe e f es r e*~ aes Ys i an” Map 1. Diburion a alle ections oor Bidder esaruined (not incase culti- vated material). Since examination of herbarium material and of living plants both in cul- tivation and in the field has failed to uncover any reliable means of separating the three previously distinguished species, we consider the genus to be mono- typic, consisting of the somewhat variable P. baccatum (L.) Krug & Urban. Other authors (Record & Hess, 1943; Adams, 1972) have expressed doubts about the distinctness of these entities, and aoe status has been advo- cated by Gillis (1974) and Correll (pers. co Although adult phyllotaxy is pea eetes alteniate: the first pair of leaves above the cotyledons on seedlings is opposite. These first true leaves are oth- erwise similar to later-formed foliage. Of the natural populations of Picroden- dron studied, all appear to be strictly dioecious. However, one tree cultivated on the estate of Mrs. Alvin R. Jennings in Coral Gables, Florida, is consistently monoecious, producing staminate flowers before pistillate ones. Other trees, presumably from the same introduction, are strictly unisexual (staminate). Thus, this tree appears to be an exceptional individual, not representing any deviant population. Of 220 herbarium specimens examined, only six were of pistillate material in flower. Pistillate flowers are probably seldom gathered because they generally appear for only a week to ten days each year, usually during the first two weeks in May, and they are green much like the leaves and are therefore inconspicuous. A profusion of common names suggests that Picrodendron 1s well known to local inhabitants throughout its geographic range. In the Bahamas it is known as blackwood or olive; in the Cayman a hae black ironwood, cherry, or wild plum; in Cuba, aceituna, gua negro, , llana, Hlanilla, mangle negro, ro- blecillo, vanilla-prieta, yana prieta, rae or yanilla-prieta; in the Dominican Lz JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Figure 2. Picrodendron baccatum: A, twig with staminate inflorescences, x .45; twig with pistillate inflorescences, x .45; C-E, staminate flowers, = 8; F, pollen se 460: G, pistillate flower, x 2.3; H, pistillate re longitudinal section, showing 2 locules (note glands at base of perianth), x 5.75: 1, ovary, longitudinal section, showing 1984] HAYDEN ET AL., PICRODENDRON 113 Republic, lemba, manzanilla (manzanillo), or mata becerro; in Haiti, gris-gris noir or simarouba; and in Jamaica, Jamaican walnut or wild plum. EXCLUDED species. Picrodendron arboreum (Miller) Planchon, London J. Bot. 5: 579. 1846, based on Toxicodendron arboreum Miller, Gard. Dict. ed. 8. Toxicodendron no. 8. 1768. Type: Miller s.n. (BM!). Rhus arborea (Miller) DC. Prodr. 2: 73. 1825, non R. arborea Macfad. Fl. Jamaica 1: 227. 1837 (= Allophylus cobbe (L.) Raeuschel, sensu Leenhouts, 1967). BIOGEOGRAPHY AND ECOLOGY Picrodendron, together with 20 other genera, has a Greater Antillean distri- bution that omits Puerto Rico (Howard, 1973). Picrodendron baccatum is one of several species (e.g., Catalpa punctata Griseb., Pseudocarpidium wrightii Millsp., Heliotropium ternatum Vahl, Linociera ee Griseb.) that are found in both the Bahamas and the Greater Antille In the Bahamas, Picrodendron grows on the ener situated on the Great and Little Bahama banks. Neither the Great nor the Little Bahama Bank has ever been connected by land to the Greater Antilles. However, during low- water stages of the Pleistocene, the greatly enlarged above-water portion of the Great Bahama Bank extended to within only 25-35 miles of the enlarged Cuban platform, greatly enhancing the chances of dispersal to the Bahamas from Cuba. Even today, the northern fringing Cuban Islands are less than 150 miles from South Andros on the Great Bahama Bank. The Great and Little Bahama banks are also believed never to have been connected to each other by land. The Northwest Providence Channel, which separates them today, has probably not been less than 20 miles wide during the last million years. Migration from Cuba to the Great Bahama Bank and thence to the Little Bahama Bank 1s not difficult to postulate. As yet, Picro- dendron has not moved across the Crooked Island Passage—a very important biogeographical barrier to plant and animal migration—to the islands of the Bahamas or the Turks and Caicos islands to the southeast. Ecologically, Picrodendron 1s a calciphile that apparently has some degree of salt tolerance. Specimens have often been collected from limestone sub- strates, either in arid habitats or near the landward margins of mangrove formations. Seifriz (1943) noted that it grows together with mangroves in Cuba, where it is often situated between zones of Avicennia germinans (L.) L. and Laguncularia racemosa (L.) Gaertner f. Howard (1955) also recorded it on Beata Island, near mangroves and in association with other coastal species such as Suriana maritima L., Thespesia populnea (L.) Solander ex Corréa, and x 11.5; J, fruit, x .75; K, 1- — paired ovules of | locule (ob = obturator, m = micropyle). (Reproduced from W. Fawcett & A. B. RENDLE, Flora of Jamaica 4: fig. 90. 1920, with permission of Trustees of British Museum (Natural History).) 114 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Ampris elemifera L. On the other hand, Asprey and Robbins (1953) included it as a constituent of ‘‘dry limestone scrub forests” in Jamaica. Presumably, suitable habitats for Picrodendron are widespread in the West Indies; its some- what restricted geographic distribution is thus all the more interesting. POLLEN PICRODENDRON (FiGuRES 3-9). The pollen of Picrodendron baccatum was de- scribed by Erdtman (1952) as 5- to 8-colporate, oblate-spheroidal, with an average diameter of 34 um, spinuliferous, with sexine thicker than nexine, and with ectosexine thicker than endosexine. We have refined this brightfheld mi- croscope description based on whole mounts and epoxy sections stained in basic fuchsin: Pollen grains 5- to 8-zonicolporate, radially symmetrical, iso- polar, oblate-spheroidal, 26 x 29 um. Amb circular. Colpi short, L: W ratio 4:1 or less, ora lalongate to circular. NPC: 545, 645, 745, 845. Exine spinose, tectate-perforate. Ectosexine about twice as thick as endosexine. Nexine ex- tremely thin and discontinuous. Electron microscope (SEM and TEM) observations add substantially to the understanding of Picrodendron pollen: Exine sculpturing echinate, the spinules to 3 um long, well spaced, with clusters of 3 to 5 around colpi, the interspinule area densely verrucate. Ectosexine 0.6-0.8 «um thick, structured from more or less cylindrical rods, with bacula incompletely fused distally, the tectal surface verrucate, traversed to varying degrees by irregular channels, some of which coincident with gaps in endosexine and nexine to form intact passage to surface. Endosexine 0.2-0.4 um thick, tapering to extinction near aperture, with narrow to broad columnar struts unevenly spaced, fused in massive columns in few areas. absent in other areas so nexine only loosely attached. Nexine extremely thin, 0.06-0.25 um, serving as footlayer for endosexinous struts, frequently discontinuous in mesocolpial regions, forming pronounced wedge-shaped thickenings of 2 distinct layers near aperture: nexine-1 (footlayer in contact with endosexine), with point of thickening initiated about | «m from colpus rim, becoming lamellate at edge of os, extending across aperture in both ace- tolyzed and unacetolyzed grains; nexine-2, with thickening moderate, initiated about | um from edge of os, appearing to extend across aperture in acetolyzed and unacetolyzed grains to form thin membrane. Intine as thick as or thicker than exine, 1-2 pm, divisible into 2 layers, outer one less electron dense, inner one granular. Intine relatively thin in mesocolpial regions, normally abutting nexine, but extending through gaps to contact sexine directly, expanding into lens-shaped thickening in aperture region and protruding up through os. OLDFIELDIOIDEAE (FIGURES 10-20). Erdtman (1952) first surveyed several mem- bers of the subfamily, but Kohler’s (1965) treatment of the pollen morphology of the biovulate Euphorbiaceae is quite comprehensive and serves in a sub- stantial way to document the brightfield microscope observations. The chief difference between Kohler’s description and our study is reflected in the NPC formulae for the group: we interpret the grains as brevicolporate instead of 1984] HAYDEN ET AL., PICRODENDRON 1s # ‘ le [te € Ficures 3-6. Pollen of Picrodendron baccatum. 3, Harris 12516, scanning electron micrograph of acetolyzed grain showing 2 slightly elongate pores, x 1800. 4-6, trans- mission electron micrographs of Clemente 6587: 4, acetolyzed grain, x 4000; 5, glutar- aldehyde-OsO,-fixed grain, x 2200: 6, glutaraldehyde-OsO,-fixed grain, x 10,000. 116 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Ficures 7-10. Brightfield photomicrographs of pollen, x 1480. 7-9, Picrodendron baccatum (Clemente 6587), single acetolyzed grain (whole-mount): 7, top focus, od in 2 short colpi; 8, optical cross section; 9, lower focus showing outline of 2 Austrobuxus carunculatus (Harlimann oo acetolyzed grain at optical cross ee nexine thickenings prominent at apert simply colpate. The apertures, whether elongate or not, are all distinctly com- pound, and we see no reason not to consider the nexine opening in each aperture as an os. Kohler refers to the os in his general description of his “7 efracoccus- type” of pollen, but he calls it an “oroid” in his “Longetia-type.” Based on our generalized brighthleld microscope observations of the taxa listed above, oldfieldioid pollen can be described as follows: 4-7 zonipororate 1984] HAYDEN ET AL., PICRODENDRON 117 Figures 11-14. Pollen of Austrobuxus carunc om 11, Hirlimann 571, scanning electron micrograph of acetolyzed grain, x 1720. 12-14, Stauffer & Hea Bye Sean electron micrographs of orains fixed in 7 lutaraldehyde-OsO,: x 6000; 1800; 14, x 6000. to colpororate, radially symmetrical. isopolar or paraisopolar, oblate-spheroidal to suboblate, medium size (25-40 um). Amb circular. Colpi lolongate, very short and inconspicuous (L:W ratio 3:1 or less), both colp1 and pores often indistinct. Ora usually larger than colpi, lalongate, round, or lolongate, delim- 118 JOURNAL OF THE ARNOLD ARBORETUM [VoOL. 65 Figures 15-20. Scanning electron oo oF neste ai Oa grains of species - Euphorbiaceae subfam. Oldfieldioideae: is (Munz 12614), x 1360: ~ Oldfeldia africana (Voorhoeve 1963 : x 1360: 17, Tetracoccus halli (Kearney & ae 7530), x 1360: 18, Piranhea longepedunculata (Blanco 299), x 1360: 19, Aus- trobuxus eugentifolius (Guillaumin & Baumann-Bodenheim 12908), x 1560: 20, same collection as in Figure 19 but different grain, showing detail of aperture. x 4080 1984] HAYDEN ET AL., PICRODENDRON 119 ited and more or less ringed by nexine thickenings formed by several more or less contiguous granules, these thickenings confined to polar ends of colpi in some species, to continuous ring or annulus around entire circumference in others. NPC: 445, 545, 645, 745. Exine tectate-perforate. Sexine much thicker than nexine, with broad-based, solid spines or spinules distributed more or less evenly over verrucate tectum except for clusters around apertures. En- dosexine baculate, bacula ramifying distally and fusing to form incomplete tectum. Nexine very thin and occasionally discontinuous, but with prominent thickenings around apertures. A limited sampling of five genera with SEM and one sample of Austrobuxus carunculatus with TEM yields the following ultrastructural description of Old- fieldioideae pollen: Exine sculpturing echinate, the spinules prevalent, becom- ing spines to 4.1 wm in Austrobuxus carunculatus, generally sharp pointed (rounded in some specimens of Tetracoccus dioicus), well spaced, the inter- spinule area sparsely (Mischodon zeylanicus) to densely verrucate. Ectosexine 0.9-1.4 um thick, structured from more or less cylindrical rods, these irregularly fused to form relatively porous, bumpy tectal surface (verrucate) traversed to varying degrees by numerous channels (perforate). Endosexine 0.1-0.4 um thick, terminating near aperture rim, with narrow to broad columnar struts unevenly spaced so nexine appears loosely attached. Nexine extremely thin, 0.06—-0.12 um, serving as footlayer for endosexinous struts, mostly continuous in interaperture region, forming pronounced wedge-shaped thickening of 2 distinct layers near aperture: nexine-! (or footlayer in contact with endosexine), with point of thickening initiated ca. | um from edge of os, appearing to extend across aperture in unacetolyzed grains as uniformly thin layer; nexine-2, with thickening initiated ca. | »m from edge of inner aperture, apparently termi- nating abruptly to form os. Intine as thick as or thicker than exine, 1-2 wm, divisible into 2 layers as in Picrodendron (inner, electron-dense area much thinner in intercolpial regions than in Picrodendron, outer, in contact with nexine, generally thick throughout grain and sprinkled with electron-dense granules and short rods). DISCUSSION OF RELATIONSHIPS RELATIONSHIPS AT THE FAMILIAL LEVEL Picrodendron conforms more closely with the Euphorbiaceae than with any other family to which relationships have been proposed—a conclusion sup- ported by gross morphology, pollen, and chromosome number (discussed be- low), as well as by vegetative anatomy (Hayden, 1977). The reduced unisexual flowers of Picrodendron fit easily within the enormous range of floral diversity exhibited by the Euphorbiaceae. The strength of this association, however, is in the presence of pendulous anatropous ovules with a ventral raphe and a micropylar obturator (FiGurE 2, I), characters diagnostic for the Euphorbiaceae. Fawcett and Rendle (1917) first pointed out these facts, but their conclusion of euphorbiaceous relationships for Picrodendron received delayed recognition because of their erroneous description of the ovary as 120 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 inferior. Although this mistake was soon corrected (Fawcett & Rendle, 1920), the damage had already been done: Pax and Hoffmann (1931) excluded Pic- rodendron from the Euphorbiaceae largely because they repeated Fawcett and Rendle’s inaccurate perception of ovary position. This unwarranted exclusion from the family by such prominent authorities discouraged any serious recon- sideration of relationships with the Euphorbiaceae for several decades. The absence of floral discs and of endosperm in mature seeds 1s unusual for the Euphorbiaceae, but these conditions are not unknown in the family. Also, Picrodendron 1s unusual (but not alone) within the Euphorbiaceae in having a drupaceous fruit instead of the more common schizocarpous capsule. These divergences are few; far more characteristics would have to be reconciled if Picrodendron were to be placed in any of the other families to which it has been assigned in the past (see Hayden, 1977, for detailed comparisons with some other families). Erdtman’s (1952) comment on the similarity of Picrodendron pollen with that of Pseudanthus Sieber ex Sprengel is largely responsible for revitalizing the notion of a relationship with the Euphorbiaceae. Indeed, the obvious sim- ilarities in pollen wall sculpturing and architecture between Picrodendron and certain Euphorbiaceae argue for a close relationship. On the other hand, Pic- rodendron shares few pollen features with the Juglandaceae (Stone & Broome, 1975) or the Rhoipteleaceae (Stone & Broome, 1971), with which it has been associated (Cronquist, 1968; Hutchinson, 1973), and it 1s not similar to the Simaroubaceae (Erdtman, 1952). Of particular note are the granular elements in the Juglandaceae (‘structure grenue”’ of Van Campo & Lugardon, 1973) that are fused in varying degrees to form the columellae of the endosexine region. The bacula of Picrodendron and Austrobuxus Miq. are typical cylindrical rods; they may vary in size but would never be characterized as granular. The per- forate ectosexine of Picrodendron and Austrobuxus is in marked contrast to the homogeneous tectum of the Juglandaceae, which is without pits or pockets except for the ultrafine channels that traverse the wall (Stone ef al., 1964). These aspects, as well as the more obvious differences in pollen sculpturing and aperture construction, leave little doubt that Picrodendron has more in common with the Euphorbiaceae than it does with the Juglandaceae. Overall, pollen of Picrodendron falls easily within Koéhler’s (1965) “*Tetracoccus-type”’ of Euphorbiaceae subfam. Oldfieldioideae. Relationship with the Oldfieldioideae is also confirmed cytologically. Fritsch (1972) reported a chromosome number of 2” = 48 for Picrodendron. Members of the Euphorbiaceae-Oldfieldioideae are not well known cytologically, but published counts of » = 24 for Mischodon Thwaites and 2n = 24 for Tetra- coccus Engelm. ex Parry (Hans, 1973) suggest that the count for Picrodendron is consistent with a base number of 12 for the subfamily. On the other hand, members of the Juglandaceae—with the exception of several tetraploid hick- ories in Carva Nutt. sect. CARYA—are uniformly 7 = 16 (Stone & Broome, 1975). The Simaroubaceae have an assortment of chromosome numbers: re- ports of » = 12 are known for Brucea J. F. Miller and Picrasma Blume, n= 13 for Castela Turpin, 2n = 26 for Holacantha A. Gray, n = 16 for Simarouba Aublet, 2” = 26 for Klainedoxa Pierre, 2m = 28 for Jrvingia Hooker f., 2n = 1984] HAYDEN ET AL., PICRODENDRON 121 36 for Quassia L., and n = 31, 2n = 62, 86 for Ai/anthus Desf. (Darlington & Wylie, 1955; Moore, 1973, 1977). Of these simaroubaceous genera, only K/ai- nedoxa and Irvingia were ever closely associated with Picrodendron (Hayden, 1977), but the known chromosome counts do not support such a relationship. The paucity of fossils in most angiosperm groups tends to confer special significance upon any known fossils. Thus, the discovery of fossilized seeds and fruits of Rosenkrantzia picrodendroides B. Koch from the Cretaceous- Tertiary boundary of West Greenland ts of interest because of their resemblance to Picrodendron (Koch, 1972). The fossil fruits are interpreted as either nuts or drupes and contain one or two seeds encased in a tough, woody pericarp (mesocarp?). The seeds are pendulous from an apical placenta and contain folded cotyledons but no endosperm. Presence of long-petiolate, trifoliolate leaves in associated fossil beds suggests an even greater list of similarities with Picrodendron. Although the fossil is unicarpellate and Picrodendron is bicar- pellate, Koch was not deterred from perceiving a close relationship between these plants; furthermore, this disparity prompted an elaborate hypothetical derivation of both plants, consistent with Small’s (1917) placement of the Pic- rodendraceae between the Juglandaceae and the Fagaceae. och’s thesis is difficult to evaluate, partly because of its complexity but mainly because of its awkward use of several morphological terms. For ex- ample, the pedicel is consistently referred to as a petiole; possible dehiscence mechanisms are discussed at some length, yet the fruit of Rosenkrantzia is described as a nut or drupe; the unicarpellate fruit is interpreted as having four dehiscence valves; and a protuberance on the fossil seed is described as a style base. Koch’s hypothetical derivation involves the fusion of paired, reflexed, unicarpellate flowers such that basal placentation (as in the Juglandaceae) gave rise to the apical placentation of Picrodendron. At the gross morphological level h’ senkrantzia and Picrodendron are merely superficial; we feel compelled to observe that such a conclusion seriously erodes the original basis for Koch’s own hypothesis Consequently, the existence of Rosenkrantzia does not strengthen the as- sociation of Picrodendron with the Juglandaceae, and we are disinclined to perceive any relationship between Rosenkrantzia and Picrodendron, although a reevaluation of this conclusion may be necessary if the four obscure valves of Rosenkrantzia prove to be evidence for a multicarpellate condition. Our proposed relationship with the Euphorbiaceae requires no elaborate derivation and is thus simpler than Koch’s hypothesis. RELATIONSHIPS WITHIN EUPHORBIACEAE The presence of two ovules per carpel and spiny pollen grains indicates that the proper placement of Picrodendron is within subfamily Oldfieldioideae (Webster, 1967). In a conspectus of suprageneric taxa of the Euphorbiaceae, Webster (1975) assigned Picrodendron to a monogeneric tribe within this subfamily. 122 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 In searching for the nearest relative of Picrodendron within the Oldfieldioi- deae, we are impressed with certain similarities to the tribe Hyaenancheae (Baillon ex Mueller-Arg.) Hutchinson, especially to members of subtribe Pai- vaeusinae Pax & Hoffm. According to Webster’s (1975) classification, the Pai- vaeusinae includes 4ristogeitonia Prain and Oldfieldia Bentham, from Africa: Celaenodendron Standley, from Mexico; and Piranhea Baillon, from northern South America. Airy Shaw (1966, 1973) has associated these genera with Pic- rodendron. Not only do they have compound leaves similar to those of Pic- rodendron, but they are also closest anatomically (Hayden, 1977, 1980) and palynologically. Webster and Lynch (unpubl.), in a more comprehensive survey of Oldfieldioideae pollen than is presented here, reached the same conclusion concerning palynological similarity of Picrodendron and the Paivaeusinae. On the basis of a combination of several morphological characters, especially the alternate, stipulate, strictly trifoliolate leaves, dioecy, and elongate styles, Pic- rodendron appears closer to the South American genus Piranhea than it does to the African Aristogeitonia or Oldfieldia. Some features of leaf anatomy of Picrodendron likewise compare most closely with Piranhea, but certain wood features are better matched by Oldfieldia (Hayden, 1977, 1980). In view of the obviously reticulate nature of relationships in the Oldfieldioideae, this does not appear to be a very serious discrepancy. Despite the above-mentioned similarities with the Paivaeusinae, Picroden- dron differs from all of the genera of Hyaenancheae in having a distinctive cluster of reproductive characters: staminate flowers with reduced calyx, borne in aments: floral disc absent in both sexes; styles elongate and basally connate: and drupaceous fruits with large, ecarunculate seeds containing plicate coty- ledons. On morphological grounds, therefore, Picrodendron appears sufhciently divergent within subfam. Oldfieldioideae to be recognized on its own. This reasoning led Webster (1975) to establish the tribe Picrodendreae (Small) Web- ster. However, when the Oldfieldioideae as a whole are better known, the strength of evidence from anatomy and palynology may prove sufficient to classify Picrodendron as an advanced subtribe adjacent to the Paivaeusinae within tribe Hyaenancheae. It may be speculated that the distinctive reproductive characters of Picro- dendron reflect an adaptive shift from insect to wind pollination. However, the lack of reduction of spines in the pollen is rather curious, since it 1s contrary to the trend evident in other taxa of anemophilous Euphorbiaceae (e.g., Hyae- nancheae subtribe Dissiliariinae Pax & Hoffm., or many genera of tribe Aca- lypheae Dumort.). Possibly there is some sort of balance between wind and insect pollination in Picrodendron, a hypothesis that needs testing by field observation. ACKNOWLEDGMENTS Although no formal grant covered this study, certain gifts, grants, and other assistance were involved in completion of the work, and we wish to acknowl- edge these with appreciation. Much of Gillis’s contribution was made while he was working under the terms of a generous anonymous grant made to the 1984] HAYDEN ET AL., PICRODENDRON 3 Arnold Arboretum of Harvard University for a revision of the Bahama flora. Some of his fieldwork was underwritten by a grant from the National Geo- graphic Society for studies in the southern Bahama Archipelago. Gillis’s notes indicated that he also wished to thank Mr. George Avery, Dr. Richard A. Howard, Mrs. Alvin R. Jennings, Dr. José de Js. Jiménez, Mr. Oris Russel, Dr. William L. Stern, and the directors of the Fairchild Tropical Garden and the U.S.D.A. Subtropical Horticulture Research Station in Miami, Florida, for advice and assistance in collecting specimens. We are grateful to the directors and curators of several herbaria (A, AAH, BM, C, F, FTG, G, GH, J, NY, P, PH, S, us, and the personal herbaria of C. D. Adams and J. J. Jiménez) for access to or loans of Picrodendron specimens. Palynological studies were supported in part by National Science Foundation grants DEB 7412661 and DEB 7724612 to D. E. Stone. Dr. Wilton R. Tenney provided photographic assistance. We thank Dr. James L. Reveal for making perceptive comments on an early draft ofthe manuscript. Publication was supported in part by a grant from the Faculty Research Committee, University of Richmond. LITERATURE CITED ADAMS, C, Z 1972. Flowering plants of Jamaica. University of the West Indies, Mona, Jamaic Airy SHAW, YL K. 1966. A dictionary of the flowering plants and ferns. ed. 7. University Press, Cambridge, England. 1973 ed. 8. University Press, Cambridge, England. ASPREY, G. F., & R. G. Rosains. 1953. The vegetation of Jamaica. Ecol. Monogr. 23: 359- os Barker, H. D., & W. S. Darpeau. 1930. Flore d’Haiti. Service technique du Dé- ae ae lAgriculture et de Enseignement gee vo rt-au-Prince. BENTHAM, G., & J. D. Hooker. 1862. Simaroubaceae. Gen. PI. Britton, N. L. 1906. Contributions to the flora of the Bahama Ge HI. Bull. New York ite 4: 137-143. Sue P. 1756. The civil and natural history of Jamaica. Printed for the author, London. ene in 1972 by Arno Press, New York. Campo, M. van, & B. LUGARDON. 1973. Structure grenue infratectale de Pectexine des Catessy, M. 1734. The natural history of Carolina, Florida, and the Bahama Islands. Vol. 2. Published by the author, London Cronauist, A. 1968. The evolution and classification of flowering plants. Houghton ifflin Co., Boston. . 1978. Ibid. Reprinted ( Ey Deets Pe q tr the author by Allen Press, Lawrence, Kansas. 1981. An integrated system of classification of flowering plants. Columbia University Press, New York. — C. D., & A. P. Marg 1955. Chromosome atlas of flowering plants. ed. 2. University Press, Aberdee ErptMAN, G. 1952. Pollen oe ke and taxonomy. Angiosperms. The Chro- nica Botanica Co., Waltham, Massachusetts Fawcett, W., & A. B. es 1917. Notes on uaa plants. J. Bot. 55: 268-271. & 1. Jamaica 4: 273-27 Fritscu, R. 1972. Ce von ee der Insel Kuba. I]. Kulturpflanze 19; 305-313. 124 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 — J.. & L. J. K. Brace. 1889. Provisional list of the plants of the Bahama slands. pa with notes ey additions by C. 8. Dottey, M.D.) Proc. Acad. oe Sci. Philadelphia 41: 349-42 Gituis, W. T. 1974. Name changes ‘a seed plants in the Bahama flora. Rhodora 76: 67-138. GOMEZ DE LA nls M., & J. T. Roic y Mesa. 1916. Flora de Cuba. Rambla, Bouza y a GRISEBACH, A ie R 1859. Flora of the British West Indian Islands. L. Reeve & Co., Londo , 1866. Catalogus plantarum Cubensium. Wilhelm Engelmann, Lei — tieR, H. 1908. On the origin of angiosperms. Bot. Gaz. (Crawfordsville) 45: 196- 98. _ 1921. Beitriige zur Kenntnis der Linaceae (DC. 1819) Dumort. 10. Die Irving- iaceen. Beih. Bot. Centralbl. 39(Abt. 2): 62-68. Hans, A. S. 1973. Chromosomal conspectus of the Euphorbiaceae. Taxon 22: 591- Haypen, W. J. 1977. Comparative anatomy and systematics of Picrodendron, genus incertae sedis. J. Arnold Arbor. 58: 257-27 1980. Systematic anatomy of Oldfieldioideae (Euphorbiaceae). Unpubl. Ph.D. dissertation, Univ. Maryland, College ses & J. L. REVEAL. 1980. Proposal for the conservation of the generic name Picrodendron Grisebach (1859) against ee yar Planchon (1846) (Euphorbi- aceae). Taxon 29: 507-511. Howarp, R. A. 1955. The vegetation of Beata and Alta Vela islands, Hispaniola. J. — Arbor. 36: 209-239 The vegetation of the Antilles. Pp. 1-38 in A. GRAHAM, ed., Vegetation and Soe history of northern Latin America. Elsevier Publ. Co., Amsterdam. Hutcuinson, J. 1973. The families of flowering plants. ed. 3. Clarendon Press, Oxford. Kocn, B. E. 1972. Fossil picrodendroid fruit from the Upper Danian of Nigssaug, West Greenland. Meddel. Gronland 193: 1-32. KOuLer, E. 1965. Die Pollenmorphologie der biovulaten Euphorbiaceae und thre Be- deutung fiir die Taxonomie. Grana Palynol. 6: 26-120. Leennouts, P. W. 1967. A conspectus of the genus 4/lophylus. Blumea 15: 301-358. Leon, H., & H. AL AIN. 1951, oe ora de Cuba. Vol. 2. Contr. Ocas. Mus. Hist. Nat. Colegio ‘De La Salle” 10: 1-45 Linnaeus, C. 1759. Systema aren ed. 10. Vol. 2. Laurentii Salvi, Stockholm. MacraDyYEN, J. 1837. The flora of Jamaica. De Orme, Brown, Green, & Long- mans, London. Ne R. J..ed. 1973. Index to plant chromosome numbers, 1967-1971. Reg. Veg. -539. ed. 1977. Index to plant chromosome numbers for 1973-1974. bid. 96: 1- DOT. Moscoso, R. M. 1943. Catalogus florae Domingensis. L. & S. Printing Co., New York. NovAk, F.A. 1961. Vy88i rostliny. Nakladatelstvi Ceskoslovenské akademie véd, Prague. Pax, F., & K. HorrMANN. 1931. Euphorbiaceae. Jn: A. ENGLER & K. PRANTL, Nat. Pflanzenfam. ed. 2. 19c: 11-253. PLANCHON, J. E. 1846. Revue de la famille des Simaroubées. London J. Bot. 5: 560- 580 Recorp, S. J., & R. W. Hess. 1943. Timbers of the New World. Yale University Press, New Haven RicHarp, A. 1845. Botanique. Plantes vasculaires. (Essai d’une flore de Pile de Cuba, Vol. 1.) Vol. 10 in R. pe LA SAGRA, Histoire physique, politique, et naturelle de Vile de Cuba. Arthus Bertrand, Paris. 1984] HAYDEN ET AL., PICRODENDRON 125 Roic y Mesa, J. T. 1945. coe cia aromaticas 0 venenosas de Cuba. Ministerio de Agricultura, Hav Sawyer, W. H. 1955. Medicinal u uses - plants by native Inaguans. Sci. Monthly 80: 371-376. ScHoLz, H. 1964. Rutales. 7a: H. Mevcuior, ed., A. Engler’s Syllabus der Pflanzen- milien. ed. 12. Band 2. Angiospermen. Gebriider Borntraeger, Berlin-Nikolassee. Serpriz, W. 1943. Plant life of Cuba. Ecol. Monogr. 13: 375-426. SLOANE, H. 1696. Catalogus pear iy quae in insula Jamaica sponte proveniunt. D. eee London. 725. A voyage to the islands Madera, Barbados, Nieves, S. Christophers and ras Printed for the author by the British Museum, London SMALL, J. K. 1917. The Jamaica walnut. J. New York Bot. Gard. 18: 180-186. Stone, D. E., & C. R. BRoome. 1971. Pollen ultrastructure: evidence for relationship of the Juglandaceae and the Rhoipteleaceae. Pollen & Spores 13: 5-14. & —— 75. Juglandaceae A. Rich. ex Kunth. World Pollen & Spore FI. 4: 1-35. ——, J. Reicu, & S. Wuitrietp. 1964. Fine structure of the walls of Juglans and Carya Site: Pollen & Spores 6: 379-392. TAKHTAJAN, A. 1966. Systema et phylogenia magnoliophytorum. “Nauka,” Moscow & Leningrad. . 1969. Flowering plants: origin and dispersal. (Transl. by C. Jerrrey.) Smith- sonian Institution Press, Washington, D. 1980. Outline of the classification of flowering plants (Magnoliophyta). Bot. v. 46: 225-359. ae R. F. 1976. A phylogenetic classification of the angiosperms. Evol. Biol. 9: Ursan, I. 1893. Additamenta ad cognitionem florae Indie occidentalis. Particula 1. Bot. Jahrb. Syst. 15: 286-361. . 1920. Simaroubaceae. Florae Domingensis. Symb. Antill. 8: 325-328. Wenster, G. L. 1967. The Sure of Euphorbiaceae in the southeastern United States. J. Arnold Arbor. 48: 303-43 — . 1975. Conspectus of a new classification of the Euphorbiaceae. Taxon 24: 593- 601. APPENDIX. Picrodendron specimens examined. Bahama Islands. Great ABAco: Old Ken’s Pt., Brace 2029 (F). ANDRos: Coconut Pt., Conch Sound, Northrop & Northrop 453 (A, F, GH); Deep Creek, Brace 5198 (F, Ny); coppice near Fresh Creek, Small & Carter 8833 (F, G, p, Us); Morgan’s Bluff, low thickets inland from rocky sea cliff, Proctor 30788 (11), Driggs Hill, Popenoe s.n., May 1965 (FTG). Cat: whiteland scrub, the Bight, Britton & Millspaugh 5884 (r, Nv); Industrious Hill, side of cliffnear caves, Byrne 323 (A), Port Howe plantation house, Byrne 364 (A); ledges in open coppice N of Stevenson, Correll 46/53 (F, FG). ELEUTHERA: rocky plains, vic. of Miller’s, Britton & Millspaugh 5593 (fF, NY). EXUMA CHAIN, GREAT GUANA Cay Britton & Millspaugh 2895 (fF, GH). LONG: Clarence Town, Britton & Millspaugh 6258 (F, Us). New Provipence: Brace 477 (Ny); Paradise Is., center of island, remnant of coppice, Gillis 11991 (a, 13). Cayman Islands. GRAND CAYMAN: logwood woodland on limestone pavement, 0.5 m1 of Joe Conger, Brunt 1783 (Bm, 11); along track between Old Isaacs and Wintersland, Proctor 15248 (3m, Gu, 1); 1.5 mi WNW-NW of Brechers, S of Joe Conger, Proctor 27976 (13); E end of island on cliff, Aings 737 (em). Lirrte CAYMAN: Snipe Point, dry, rocky woodland, Proctor 28038 (BM, 11); S side of Blossom Pt., Kings 77 (BM, NY). CAYMAN Brac: W end, edge of dense woodland, Proctor 29125 (1). 126 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Swan Islands. Great Swan: lat. 17°25'N, long. 83°56'W, Proctor 32534 (BM, 11). Cuba. Camactiey: Silla de Cayo, Cayo Romano, Shafer 2505 ~ F, GH, NY, P, us); Santa Lucia, Shafer 965 (Nv); Cayo Sabinal, Shafer 1107 (BM, F, NY, US). HABANA Marianao Beach, coastal thickets, Le6n s.n., Feb. 1921 (Gu); Pr. ae. Ekman 1207 (G, S, US). ORIENTE: Sardinero, coastal thickets, Clemente 6587 (Gu, Us); Manigua Costera, Plaza de Puerto Padre, Lopez 26 (us); Guantanamo Bay, U.S. Naval Station, Britton 2259 (Ny); Santiago de Cuba, Sa ee wayside coastal thickets, Clemente 6604 (Gu); Santiago de Cuba, Juragria Beach, coastal cliff, Clemente 7436 (GH, us); Santiago de Cuba, Siboney, in calcareous soil, EAman 8710 (F, us); Santiago de Cuba, near El Nacro along seacoast, Ekman 9195 (F, G, us); Banes, Puerto Rico, coastal forest, Ekman 6625 (F, G, Us); San German, along railroad in forest, Ekman 6344 (G); Cabo Cruz, coastal eens Leon 16351 (Gu, J, Us); Punta Piedra, Nipe Bay, Britton et al. 12460 (F, Gu, , US). PINAR DEL Rio: Bahia Honda, Finca Toscano, coastal thickets, Alain 1803 (GH, WJ, + Las VILLAs: Cienfuegos, Caunao R., Iguana Point, Jack 5/26 (aA, Pp, Us); Caunao R. to Cienfuegos Bay, Jack 75/6 (F, us); Cienfuegos Bay, Punta Sabanilla, coastal thicket, Britton et al. 4584 (Ny); Punta Diablo, Cienfuegos Bay, Britton & Wilson 5672 (F); Cienfuegos, Faro Villa Nueva, hills and cliffs, Castillo de Jagua, Combs 646 (Gu); Camino de la Costa, E of Castillo de Jagua, Howard et al. 249 (a, Ny); Soledad, Gavilan, Jack 5824 (A, p, US); Gavilan, Rehder 1102 (a, pv), along Arimao R., Bangham s.n., 22 July 1929 (J). WITHOUT DEFINITE LOCALITY: Sagra s.n. (vp, Herb. Richard) Dominican Republic. Prov. AzUA: Monte Rio, Lavastrée 1750 (ny), NE of Azua, Rose et al. 4042 (ny, US). Prov. BARAHONA: near Barahona, Fuertes 1535 (BM, F, GH, NY, P. us); Barahona, Jiménez s.n. (NY); Beata Is., edge of lagoon on E side, Howard | aaa BM, Us); Beata Is., without further locality, Ostenfeld 297 (c). Prov. LA Vea: La Veg City, eases La Escuela de Agricultura, Padre Cicero et al. 7020 (A, 1), ran 7125 (a, F, 11); Bayacanes, near La Vega City. serpentine hills, 300-350 m, Liogier 11927 (Fo, Ny), Prov. Monte Cristi El Tamarindo, Hatillo Palma, Marcano 5202 (Ny, Herb. Jiménez). Prov. PEDERNALES: SE of Cabo Rojo near beach in dogtooth limestone, Liogier 13910 (Ny); Cabo Rojo in thickets, sea level, Liogier 16872 (Ny), Cabo Rojo, eee 4528 (Herb. Jiménez). Prov. Et Seiso: E of Jovero, between Punta Jiaco Punta Rey, Abbott 2872 (us). Haiti. [ue pe LA GonAve: Mahautiére, edge of mangrove, Ekman 8685 (c, Gu); La Mahautiére, Eyerdam &4 (a, F, GH, US); Dubedon, 200 m, Buch 1009 (1). DEP. pe VPOuEST: Cabaret, Baie des Moustiques, Leonard & Leonard 12004 (a, Us Jamaica. CLARENDON PARISH: Jackson Bay, dry, rocky woodland near sea, Proctor 29171 (J, NY); Jackson Bay, 0.7 mi ee wee in on nd, border of salina, Proctor 32643 (F, 11); Stee Ridge, limestone near seacoast. Van der Porten s.n., 11 July 1950 (3); Portland Ridge, coastal hammock, Hes <. Arnold 148 (Gu). St. ANDREW PArisH: above Ferry Grabham s.n., 11 Jan. 1896 (Ny); Ferry Peninsula, 100 ft elev.. Campbell 6182 (Bm, F): limestone hillside near Ferry, Harris 12516 (bm, GH, NY, PH, US). ST. CATHERINE PARISH: W base of Port Henderson Hill, scrub woodland, Proctor 22129 (Gu, 11), Port Henderson Hill, Webster & Wilson 4929 (a, 11, US); summit of Port Henderson Hill, ca. 700 fi, thickets, Proctor 32808 (13); Port Henderson Hill, path to Rodney’s Lookout, Adams 12801] (sm); Bracton & Rodney’s Lookout, Gibbs s.n. (Herb. C. D. Adams); near Ft. Clarence, E side of Great Salt Pond, Lewis s.n., 1 March 1951 (11); Old House Point, on ee limestone, Du Quesnay 555 (13); Portland Ridge near coast, Howard 12001 (a, 1), is); Ferry River, Spanish Town Road, [Harris 9046 (F. NY. US): a Bay, Tullock os Great Goat Is., rocky woodland, Proctor 17 497 (A, F, Ww, US); Great Goat Is., Harris 9313 (A, BM, C, NY), Harris 12519 (3M, GH, NY, PH, US); Little ae Is., Old Harbor Bay, near mangrove swamp, Britton & Hollick 1851 (Nv). WitHour Locality: Macfadyen sm. 1984] HAYDEN ET AL., PICRODENDRON [27 Cultivated. Topaco: cultivated at Botanic Station, Broadway s.n., 8 Nov. 1932 (BM). TRINIDAD: Royal Botanic ee of-Spain, Fairchild 2046 (Aan), Nevling 316 (AAH), Broadway 7321 (A, AAH, F, _U.S. A., Froripa, Miami: U.S.D.A. Plant Intro- duction Station, P.I. 98559, M5830, introduced from Trinidad Botanic Garden, Gillis 7846 (AAH, FTG, 1), Fennell 948 (Aan); estate of Mrs. Alvin R. Jennings, Rehder s.n., 30 April 1920 (AAH), Gillis 9330 (AAH, FTG), Popenoe 20 (FTG); Fairchild Tropical Garden, introduced from Driggs Hill, South Andros, Bahama Islands, Gillis 10924, 10975, 11031 (AAH, FTG, J). C.R. B. W.J.H PLANT VARIETY PROTECTION OFFICE DEPARTMENT OF BIOLOGY BELTSVILLE AGRICULTURAL RESEARCH UNIVERSITY OF RICHMOND Center, U.S.D.A. RICHMOND, VIRGINIA 23173 BELTSVILLE, MARYLAND 20705 G.L. W. D. E..8. DEPARTMENT OF BOTANY DEPARTMENT OF BOTANY UNIVERSITY OF CALIFORNIA DuKke UNIVERSITY Davis, CALIFORNIA 95616 DuRHAM, NortH CAROLINA 27706 BURTT, STAURANTHERA 129 THE FIRST SPECIES OF STAURANTHERA (GESNERIACEAE) FROM NEW GUINEA, WITH GENERAL NOTES ON THE GENUS B. L. BurtTT ALTHOUGH Stauranthera Bentham is listed in Willis (1973) as a genus of ten species, on closer study that number proves to have been a little generous. To date, twelve names have been published. Stauranthera chiritiflora Oliver has become the type species of Whytockia W. W. Sm. (Smith, 1919), and S. tsiang- iana Hand.-Mazz. also belongs to Whytockia (Weber, 1982). Merrill (1923) reduced both S. ecalcarata R. Br. and S. philippinensis Elmer to synonymy under S. caerulea (Blume) Merr., but for the present I am inclined not to include S. philippinensis there. Stauranthera brandisii C. B. Clarke, described almost simultaneously as Rhynchotechum brandisii C. B. Clarke, 1s of uncertain identity but is not a Stauranthera (see Burtt, 1962, p. 37, to which it can now be added that on the isotype at Calcutta, infructescence and leaves are joined, so there is no doubt that they belong together). Finally, I now reduce S. jo- hannis-winkleri Kranzlin to a synonym of S. argyrescens Hallier f. This leaves seven species, ranging from northeastern India and Bangladesh to Celebes (Sulawesi): Stauranthera umbrosa (Griff.) C. B. Clarke, from NE India, Burma, S China, and the Malay Peninsula; S. grandiflora Bentham, from NE India, Bangladesh, Burma, Thailand, the Malay Peninsula, and Sumatra; S. caerulea (Blume) Merr., from Sumatra, Java, and (?) Celebes; S. tonantha Hallier f. and S. parvifolia S. Moore, both endemic to Sumatra; S. argyrescens Hallier f., from Borneo; and S. philippinensis Elmer, from the Philippines. Some of these records (partly culled from the literature) require verification. I have not seen S. jonantha or S. parvifolia. The distinction between S. umbrosa and S. caerulea is not always clear. The discovery of a distinct species in New Guinea greatly extends the generic range of Stauranthera. When a western Malesian genus is discovered in New Guinea, it is natural to seek possible lines of contact as indicated by the affinities of the species. Unfortunately, S. novoguineensis does not provide particularly strong clues, especially while the S. caerulea—S. philippinensis problem awaits clarification. There seem to be two affinities, as indicated in the diagnosis. One, with the eastern Bornean S. argyrescens Hallier f., rests on just a single character that is uniquely shared by these two species: the presence of filiform sclereids (Hallier’s ‘‘Faserzellen’’?) in the mesophyll. These may be simple or slightly branched toward the tips. They are not known in other species, and since their presence can be detected in the dried leaf by the characteristic ripples on the surface (easily visible with a x 10 hand lens), it seems fairly certain that they © President and Fellows of Harvard College, 1984. Journal of the Arnold Arborenim 65: 129-133. January, 1984. 130 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 do not occur. In general facies, especially in leaf shape, S. novoguineensis 1S closer to S. philippinensis—a species that lacks sclereids and seems to have slightly different dae but nevertheless appears to provide the closest affinity for the new spec An additional noratie link between New Guinea and the Philippines is always of interest and merits comparison with other examples within the family. In Agalmyla Blume (incl. Dichrotrichum Reinw.) the link is evidenced by the fact that most of the species with long-pedunculate inflorescences are found in New Guinea, the Moluccas, and the Philippines. Epithema benthamii C. B. Clarke, described from the Philippines, occurs also on the New Guinea mainland. Cyrtandra tarsodes B. L. Burtt, of New Guinea, is most closely related to C. auriculata C. B. Clarke, from the Philippines (Burtt, 1971). These two species are rather isolated in the genus as a whole, but in general the much-branched, shrubby New Guinea species of Cyrtandra are more closely related to the many species of similar habit in the Philippines than they are to the larger-leaved, single-stemmed species that predominate in Borneo. In a collection of essays on Wallace’s line (Whitmore, 1980), the composition of the eastern Malesian flora and fauna is reexamined in relation to the recent findings on plate tectonics. It is suggested that Australia-New Guinea drifted northward and made contact with the Laurasian outliers of Celebes about 15 million years ago. Celebes may have a central suture, making it half Asiatic and half Australasian. At about the same time, New Guinea collided with a “tertiary island” to the north, eventually resulting in the upthrusting of the central mountain spine; New Guinea is thus also of mixed “parentage.” In the botanical essays in this volume, attention is focused on Wallace’s line and on the east-west interchange through the unstable archipelagic area between Celebes and New Guinea termed ““Wallacea” by a number of dia eer there is no further mention of the “tertiary island” or “tertiary a Raven 1979) with which New Guinea is said to have collided. Ifit carried a iene type flora, it could well have been responsible for the New Guinea—Philippine links that Corner (e.g., 1967, p. 45) ascribed on phytogeographic grounds to an old Melanesian foreland. Stauranthera is a very distinctive genus marked by its strongly plicate calyx and by the cruciform arrangement of its four anthers. It is always strongly anisophyllous, with the small leaf reduced to a lateral auricle. The structure of its shoot and inflorescence has recently been elucidated by A. Weber (1977); he suggests a fairly close affinity to Loxonia Jack, with which | certainly agree. Stauranthera grandiflora Bentham, the type species of the genus, has a strongly spurred corolla—a feature included in the original generic ee but not present in any other species and clearly not of generic significan Herbarium material of Stauranthera is often inadequate: Lie fall quickly, and the somewhat succulent plants require more careful drying than they some- times receive. Although a wider range of good specimens is essential before the genus can be satisfactorily revised, the records of two species, S. argyrescens and S. grandiflora, are extended here. 1984] BURTT, STAURANTHERA 131 Stauranthera argyrescens Hallier f. Ann. Jard. Bot. Buitenzorg 13: 286. 1896. Type: Kalimantan, Amai Ambit, ca. 600 m alt., cult. in Horto Bogo- riensi, Hallier B3453 (Bo, n.v.). S. johannis-winkleri Kranzlin, Mitt. Inst. Allg. Bot. Hamburg 7: 92. 1927. Type: West Borneo [Kalim Sree Bukit Mulu, Urwald, ca. 700 m alt., | December 1924, Wink- ler 453 (HBG, photo SPECIMENS EXAMINED. Indonesia. BorNeo. Kalimantan Timur: Central Kutei, Belajan R., G. Kelepoh, near Tabang, forest, sandy yellow loam, in rivulet, 50 m alt., Kostermans 10429 (L); around Jellini, along S Belajan, NW of Tabang, tropical rain forest, 100-150 m alt., Murata et al. B1126 (1); Gunung Kongkat-G. Kongbotak, 1°10’N, 116°20'E, lowland rain forest, 150-200 m alt., Kata & Wiriadinata BS 156 (L); W Kutei, G. Kelepoh, near Tabang on Belajan R., yellow loam, low alt., Kostermans 10596A (1); E Kutei, § Menubar region, ridge, loam soil with limestone, 30 m alt., Kostermans 5228 (1). I have not seen the type of Stauranthera argyrescens, but I have little doubt that all the material cited above belongs to this species. Hallier described a plant that flowered in the Botanic Garden after having been transported from Borneo. His illustration suggests a stunted plant that had not fully recovered from the move. The other specimens all have a distinctly pedunculate inflo- rescence and obvious internodes. Hallier derived his specific epithet from the silvery patches on the leaves—a character not noted by other collectors, but one that is not constant in other rain-forest herbs (cf. Burtt, 1976). This species has distinctive filiform leaf-sclereids that were noted by Hallier and are present in all the material cited, including the type of S. johannis-winkleri. They are not known in any other species except S. novoguineensis (described below). tees grandiflora Bentham, Scroph. Ind. 57. 1835—sphalmate “‘gran- a”, C. B. Clarke in A. & C. DC. Monogr. Phanerog. 5: 190. 1883, nee in Fidoker f. Fl. Brit. India 4: 371. 1884. Type: Penang, Wallich 6395 (K). Glossanthus? grandiflorus Bentham in Wallich, Num. List, 6395. 1832, nomen nudum. GENERAL DISTRIBUTION. Thailand, Burma, Bangladesh, Malay Peninsula, Su- matra. New recorps. India. GREAT Nicosar Is.: Laful, rocky streambed inside dense forest, 10 m alt., Hore 8777 (e, pp). Malay Peninsula, SELANGOR: Ulu Gombak, descent to river at mile 22, ca. 427 m alt., Burtt & Woods B1610 (k). The species has not previously been recorded from the Nicobar Islands. The record from Ulu Gombak is new merely from a local point of view: the species is not in Henderson’s list of Kuala Lumpur plants (Henderson, 1928). This species is often referred to as Stauranthera grandifolia, as it was orig- inally printed. However, C. B. Clarke reported that Bentham himself admitted that this was a printing error, and the citation of the earlier Glossanthus? grandiflorus at the time of publication confirms this. The species should there- fore be known as S. grandiflora. 132 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Stauranthera novoguineensis B. L. Burtt, sp. nov. Stauranthera philippinensi Elmer affinis, sed ovario ellipsoideo glabro (nec sphaerico pubescente), stylo longiore, stigmate bilamellato (nec capitato), foliis sclereidis filiformibus praeditis distinguenda. S. argyrescens Hallier f., sclereidis foliorum S. novoguineensi congruit sed habitu humiliore et foliis oblique ob- ovatis apice rotundatis longe recedit. Herba caule ad 10 cm longo erecto vel basi decumbente in siccitate brunneo longitudinaliter rugoso (in vivo subcarnoso?) basi radicante. Folia ut videtur alterna, altero cujusque paris ad auriculam lateralem ca. 2 x 4 mm redacto; petiolus 1-2.5 cm longus, pilis brevibus patentibus indutus; lamina 10-1 4—6 cm longa, inaequaliter (subfalcatim) elliptica, acuta vel breviter acuminata, basi inaequilateraliter angustata, marginibus irregulariter acute dentato-serrata, supra glabra in siccitate sclereidis rugulosa, infra ad venas leviter prominentes et ad venulas pilis brevibus saepe hamatis pubescens, venis lateralibus utrin- secus 5-7. Inflorescentia terminalis, simplex vel basi ramo uno praedita, ad 10 cm longa; pedunculus 4—7 cm, dimidio superiore bracteis sterilibus |—2 ornatus, glaber; pedicelli 0.5-1 cm longi, pilis saepe hamatis breviter pilosi, ebracteati. Calyx ca. 7 mm longus, ad trientem 5-lobus, sinubus conspicue plicatus, extra pilis saepe hamatis indutus, intus brevissime pubescens pilis paucis hamatis. Corolla rotata, ca. | cm longa, ad mediam subaequaliter 5-lobata, lobis ca. 5 5 mm apice rotundatis marginibus undulatis. Stamina fertilia 4, basi corollae orientia: filamenta filiformia ca. 2 mm longa, glabra; antherae obtuse trian- gulares, basi leviter cordatae, apicibus cruciatim cohaerentes. Ovarium ovo- ideo-oblongum, 1.5 mm longum, glabrum, in stylum fere aequilongum robus- tum parce patenter pubescentem transiens; stigma bilamellatum, marginibus ciliis planis ornatis. Type. Papua New Guinea, West Sepik Prov., Carpentaria Expedition Ge Camp, stream margin from K1 to K18 helipad, 340 m alt., January 1978, S. Hoover 832 (A) AbDDITIONAL SPECIMENS. Papua New Guinea. West Sepik Prov. Telefomin Subdistr.: Prospect Creek near Frieda R., 4°42’S, 141°48’E, valley forest, 445 m alt., Henty & Foreman NGF 42601] (1); iain ea Expedition Base Camp, stream margin NE of ridge above Storm Creek, 550 m alt., Hoover 768 (A). ACKNOWLEDGMENTS I wish to thank P. F. Stevens (Arnold Arboretum) for making W. Scott Hoover’s New Guinea material available to me, and N. P. Balakrishnan (Bo- tanical Survey of India) for the specimen from the Nicobar Islands. I recently visited the Rijksherbarium, Leiden, and I am greatly indebted to the director, C. C. Kalkman, for facilities there. Anton Weber (Vienna) kindly criticized the first draft. REFERENCES Burtt, B. L. 1962. Studies in the Gesneriaceae of the Old Aa d, XXI1. Rhynchotechum and /santhera. Notes Roy. Gard. Edinburgh 24: 35 1984] BURTT, STAURANTHERA 133 1971. Studies in the ae of the Old World, XXXIV. A miscellany from south eastern Asia. [bid. 31: 35-52 1 ~ tes on rain-forest i Gard. Bull. Straits Settlem. 29: 73-80. Garner E. Je 1967. Ficus in the Solomon Islands a Jurassic ees of Melanesia. Philos. Trans., ser. B. 253: HENDERSON, M.R. 1928. The flowering plants of Kuala on in the Malay Pen- sula. Gard. Bull. Straits Settlem. 4: 24-373. Aas E. D. of Printing, Manila RAvEN, P. 1979. Plate tectonics and southern hemisphere biogeography. Pp. 3-24 in K. Larsen & L. B. Ho_m-Nie.sen, eds., Tropical botany. Academic Press, New York nd - bearing on the post- -159. The flowering plants of the Philippine Islands. Vol. 3. Bureau ork. SmitH, W. W. 1919. Whytockia, a new genus of Gesneriaceae. Trans. Bot. Soc. Edin- 339. : eitrage zur Morphologie und Systematik der Klugieae und Loxonieae (Gesneriaceae), VI. Morphologie und Verwandtschaftsbeziehungen von Loxonia und Stauranthera. Flora, ser. B. 166: 153-175. ontribution to the morphology and systematics of Klugieae and Lox- onieae (Gesneriaceae), IX. The genus Whytockia. Notes Roy. Bot. Gard. Edinburgh 40: 359-367. Wuitmore, T.C., ed. 1980. Wallace’s line and plate tectonics. Oxford University Press, Oxford. Wiis, J.C. 1973. A dictionary of the flowering plants and ferns. ed. 8 (revised by H K. Airy SHAW). Cambridge University Press, Cambridge, England. RoyaL Botanic GARDEN EpINBURGH EH3 5LR, SCOTLAND RAMAMOORTHY, SALVIA 135 NOTES ON SALVIA (LABIATAE) IN MEXICO, WITH THREE NEW SPECIES T. P. RAMAMOORTHY THE GENUS Salvia L., well known to horticulturists, is an outstanding member of the Mexican flora. With its bright red, blue, sometimes yellow, or occasion- ally white flowers, it is a very conspicuous element in the highlands of Mexico. The flowers range in length from ca. 8 mm (S. hispanica) to 6 cm (S. fulgens). Common pollinators are hummingbirds, bees, and butterflies. A very high percentage of species are perennials; there are very few annual species of Sa/via in Mexico. Standley and Williams (1973) estimated that the genus consists of over 900 species, with nearly 500 from the Americas. Fernald’s (1900) synopsis of the Mexican and Central American species of Sa/via updated Bentham’s (1848) and Hemsley’s (1882) accounts. Bentham recognized 118 species, Hemsley 126, and Fernald 217. Epling (1939), perhaps the most distinguished student of the genus, recog- nized 233 species for Mexico (all belonging to subg. CALOPHACE (Bentham) Epling). Twenty sections and 200 species are endemic to Mexico. In a subse- quent series of supplementary notes (1940, 1951, 1957, 1962), he described several more species. The increase in botanical activity in Mexico during the last few years has resulted in many new specimens that were not available to earlier botanists. I have studied the specimens of Sa/via among these collections, along with the very rich holdings of the genus in the Gray Herbarium, Harvard University. This is the first report resulting from my study. It now seems certain that over 275 species of Sa/via (30 percent of the genus) occur in Mexico; of these, 88 percent (over 26.7 percent of the genus) are endemic to Mexico. Mexico has indeed been a primary center of speciation for Salvia. Three new species are described here. A commentary on Salvia tubifera is also provided. Salvia anastomosans Ramamoorthy, sp. nov. FIGURE |. Caulis pilis ramosis adpressis gerens. Foliorum laminae |.5-3.7 cm longae, 0.5-1 cm latae, oblongae, ad extremos rotundatae, supra glabrae, subtus ad- presse albo-tomentosae; venae anastomosantes. Calyx albo-tomentosus, ca. 7.5 mm longus, ca. 3 mm latus. Corolla azurea, ca. 1.2 cm longa. Erect perennial shrub to 2 m tall, profusely branched. Stem terete, appressed © President and Fellows of Harvard College. 1984. Journal of the Arnold Arboretum 68: 135-143. January. 984. 136 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 : A basis A} Figure 1. Salvia anastomosans (from holotype): a, habit: b, leaf, c, flower; d, dissected corolla. 1984] RAMAMOORTHY, SALVIA 137 white-tomentose with branched hairs when young, becoming glabrous with age. Petiole 0.5-1 cm long, appressed white-tomentose with branched hairs: leaf blade oblong, 1.5-3.7 by 0.5-1 cm, rounded at both ends, entire, glabrous above, appressed white-tomentose beneath with branched hairs, the midrib pronounced, especially beneath, with 9 to 12 pairs of lateral veins, secondary veins numerous and anastomosing into fine network very pronounced against white pubescence. Inflorescences axillary and terminal racemes of interrupted verticils, the internodes separating verticils 0.5-1 cm long, appressed white- tomentose with branched hairs; bracts small, inconspicuous, white-tomentose, early deciduous; pedicels 1-2 mm long, white-tomentose. Calyx ca. 7.5 by 3 mm, bearing numerous sessile glands, these covered by white tomentum of branched hairs, the upper lip entire, acuminate, the lower with 2 acuminate lobes; corolla ca. 1.2 cm long (tube 6 by 3 mm), blue, the upper lip nearly entire, obscurely emarginate, bearded above, as long as lower lip, the lower lip 3-lobed, with lateral lobes shorter and rounded, and middle lobe ca. 6 by 6 mm, broadly rounded, emarginate, undulate; stamens 2, included, filaments 2 mm long, connective ca. 4 mm long, anthers ca. 2 mm long; style ca. 1.4 cm long, slightly exceeding corolla, divided into 2 unequal arms, glabrous. Type. Mexico, Oaxaca, Teposculula, above Tamazulapan (alt. 2250 m), ‘“‘ma- torral” with Bursera, tree Ipomoea, etc., Aug. 8, 1981, D. Lorence et al. 3709 (holotype, MEXU). The specific epithet refers to the impressively anastomosing venation pattern that helps to distinguish the species immediately. Sa/via anastomosans belongs to sect. TOMENTELLAE Epling and Is related to S. candicans, which differs in having truncate leaf bases and less obvious venation. Salvia boegei Ramamoorthy, sp. nov. FIGURE 2. Petioli 2-6 mm longi, prope basin articulati, albo-villosi. Foliorum laminae 1-2.8 cm longae, 0.8-1.9 cm latae, ovatae, ad apicem acutae, ad basim trun- catae vel subcordatae, supra puberulae, subtus ad nervos villosae. Verticilli fere capitati 2 cm lati. Calyx albo-lanosus. Corolla azurea, 2—2.1 cm longa, tubo papiullato. Erect perennial shrub or subshrub (height not known), profusely branched. Stem reddish, angled, white-tomentose to villous, glabrous with age. Petiole 0.2-0.6 cm long, articulated near base, white-villous; leaf blade ovate, 1-2.8 by 0.8-1.9 cm, acute at apex, truncate to subcordate at base, crenate-serrate at margin, puberulent above, appressed-villous along nerves beneath and with secondary mosing. Inflorescences terminal racemes Locman verticils, the internodes the upper lip entire, rounded, the lower obscurely 3-lobed, with lateral lobes — 38 FiGure 2. dissected corolla JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Salvia boegei (from holotype): a, habit; b, node with leaves; c, flower; d, l 1984] RAMAMOORTHY, SALVIA 139 shorter, all rounded; corolla 2—2.1 cm long (tube 6 mm long, with pair of papillae toward base on inside), blue, the upper lip 4.5 mm long, as long as lower, emarginate, bearded on back, bearing many sessile glands, these white when dry, the lower lip ca. 4.5 by 4.5 mm, villous on outside, bearing glands, 3-lobed, with lateral lobes shorter, middle lobe rounded and undulate; stamens 2, included, filaments ca. 2 mm long, connective ca. 3.5 mm long, anthers ca. 2 mm long; style 1.1-1.2 cm long, hairy, barely exserted, divided into 2 unequal arms. Type. Mexico, Puebla, Puente de Dios Molcaxac (alt. 1800 m), with Dasylirion, Agave, Nolina, Yucca, Sept. 16, 1971, W. Boege 1933 (holotype, MEXU). This species is named after Mr. Wolfgang Boege, an inspiring teacher and plant collector, whose excellent personal herbarium is now with the National Herbarium of Mexico (mExu). He died in Jalisco in 1975 while on a plant- collecting expedition. Salvia boegei can be recognized immediately by its articulated petiole, almost capitate verticils, and woolly white calyx. It belongs to sect. SCORODONIA Epling and appears to be related to S. breviflora, which differs in having tardily de- ciduous bracts and leaves that are whitish beneath. Salvia rzedowskii Ramamoorthy, sp. nov. FIGURE 3. Caulis pilis albis reflexis obsitus. Folia crassa, 2—6 cm longa, 0.6-1.6 cm lata, elliptica, ad basin et ad apicem acuta, supra pilosa, subtus albo-lanata. Verticilli congesti. Calyx villosus, ca. 1.1 cm longus, 2.5 mm latus. Corolla alba, ca. 1.3 cm longa. Erect, perennial, suffrutescent herb to 6 dm tall. branched. Stem 4-angled, sulcate, tomentose with white, reflexed hairs, pilose to glabrous with age. Petiole 1-3 mm long, villous; leaf blade thick, elliptic, 2-6 by 0.6-1.6 cm, acute at both ends, crenate-serrate at margin, pilose above, with 6 to 8 pairs of faint lateral veins, white-woolly below. Inflorescences terminal racemes of crowded eae internodes separating verticils not well pronounced; bracts small, ca. |-2 y 4 mm, ovate, caudate-acuminate; pedicels 2-4 mm long, terete, cil spreading at anthesis. Calyx villous, ca. 1.1 cm by 2.5 mm, bilabiate, the upper lip ovate, acuminate, entire, the lower with 2 ovate, acuminate lobes: corolla ca. 1.3 cm long (tube ca. 8 by 2.5 mm, glabrous), white, the upper lip 5 mm long, rounded and emarginate at tip, bearded, the lower lip ca. 6 by 6 mm, 3-lobed, with lateral lobes shorter and rounded, middle lobe undulate: stamens 2, included, filaments ca. 2.5 mm long, connective ca. 2 mm long, anther ca. 2mm long; style ca. 1.2 cm long, hairy, divided into 2 unequal arms. Type. Mexico, México, 5 km SW of Sultepec. on the road to Amatepec, me- sophyllous forests (alt. 2350 m), February 18, 1979, J. Rzedowski 36071 (ho- lotype, MEXU). The specific epithet honors Dr. Jerzy Rzedowski, whose contribution to Mexican botany is well known. Fic coro JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 suRE 3. Salvia rzedowskii (from holotype): a, habit; b, leaf; c, flower; d, dissected lla. 1984] RAMAMOORTHY, SALVIA 14] Salvia rzedowskii is easily recognized by its thick leaves that are white-woolly beneath, its condensed raceme (resembling a nearly cone-shaped head), its caudate-acuminate bracts, and its white flowers. White flowers in Sa/via are rare, and the species may be moth pollinated. The affinities of this species are not clear. SALVIA TUBIFERA, A LITTLE-KNOWN SPECIES Cavanilles (1791) described Salvia tubifera from cultivated plants grown from seeds brought from Mexico. No exact locality for the original seed col- lection 1s known. Fernald (1900) included this species in his synopsis of Mexican and Central American salvias but did not cite any additional specimens. Epling assigned it to sect. TUBIFLORAE Epling and cited two specimens, one from Real del Monte, Hidalgo, Mexico, and the other from the central valley of Mexico. I have seen both. The species has been re-collected twice from near Real del Monte, in 1950 by Matuda and in 1982 by Benitez. Recent studies have shown that the species is, in fact, widespread and has been going by the names S. excelsa and S. venosa. Salvia excelsa was described by Bentham from plants grown from seeds supposedly of Guatemalan origin. I have seen a photograph of the type, old collections in the Gray Herbarium (Gu), and recent collections in the Herbario Nacional (MExU) and the herbarium of the Escuela Nacional de Ciencias Bio- logicas (ENCB). These clearly match Cavanilles’s plate of S. tubifera Cav. and the type photo of the species. Salvia venosa, described by Fernald (1900), is based on a collection from near San Crist6bal las Casas in Chiapas, Mexico (Nelson 3138 (Gu, us)). Epling (1939) separated S. venosa and S. excelsa on geographic grounds (Chiapas vs. Guatemala) but later (1940) concluded that they were conspecific. I have seen the type material at GH and am of the same opinion. These plants that have been going by three different names represent only one species, for which the earliest name is S. tubifera Cav. Complete specimen citation with synonymy and a description are provided ere Salvia tubifera Cav. Ic. Descr. Pl. 1: 16. ¢. 25. 1791 (illustration and description of plants grown from seeds brought from Mexico). LEcToTyPE: we have recently received in MEXU photographs of types of all salvias in the collections at MA described by Cavanilles. There are two specimens labeled S. tubifera, and I designate the specimen with the annotation “\S. tubifera Icon. t. 25” as lectotype. Epling did not typify the name in 1939 pe Eg Willd. Sp. Pl. 1: 141. 1797, nom. superfl. Type: Willdenow cited a um in herb. Jussieu vidi’; this may be considered as the type of his ae Salvia pee Bentham, Bot. Gaz. 27: 90. 1841. Type: description from plants grown from seeds brought from ?Guatemala (holotype, specimen in k, n.v., photo GH!). 142 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Salvia venosa Fern. Proc. Amer. Acad. Arts 35: 545. 1900. Tyre: Mexico, Chiapas, San Cristobal las Casas, Nelson 3138 (holotype, Gu!; isotype, Us!). Herb or subshrub 0.5-1.2 dm tall, branched. Stem 4-angled, sulcate, villous when young, sparsely pilose to glabrous with age, the hairs multicellular, often crisped, usually appressed, often persisting along grooves. Petiole 2-5 cm long, villous, the hairs usually crisped: leaf blade ovate, sometimes nearly rotund- ovate, 4-12 by 2.5-7.1 cm, apex acute to acuminate, base broadly to narrowly cuneate to somewhat truncate, margin crenate-serrate, lateral veins 4 to 7 pairs, upper surface pilose to glabrescent, lower surface densely to moderately villous to nearly glabrescent and paler. Inflorescences terminal racemes of interrupted, - to 6-flowered verticils separated by internodes of 0.5-4 cm; bracts 4 mm-— 1.2 cm by 4-8 mm, broadly ovate, caudate-acuminate, pilose abaxially, early deciduous; pedicels |.5-8 mm long, angled, villous. Calyx 8 mm-I.1 cm by 3-4 mm, pilose to glabrous above, usually villous to hispid below, bilabiate, upper lip entire, lower 2-lobed, all acuminate; corolla 2.6-3 cm long (tube 2.2- 2.4 cm by 3-4 mm), scarlet, villous, the upper lip 5-6 mm long, longer than lower, rounded, emarginate, enclosing stamens and stigma, the lower lip 3-4 y 3-4 mm, 3-lobed, with lateral lobes shorter, middle lobe larger and rounded, all reflexed; stamens 2, the filaments 3 mm long, attached near base of corolla tube, the connective 2—2.2 cm long, the anthers 2.7 by 0.5 mm, acute at ends: style as long as corolla tube, pilose, ending in 2 unequal stigmatic lobes. Seeds not seen. Flowering July to November. DistRiBpUTION. Nuevo Leon in northern Mexico (Epling (1951) cited Meyer & Rogers 2891 from Nuevo Leén, but I have not seen it) through Hidalgo, Puebla, Guerrero, and western Veracruz to Chiapas and ?Guatemala in the southeast. Usually in pine/oak forests; 1800-2750 m alt. SPECIMENS EXAMINED. Mexico. HipAtco: Zimapan, between Zimapan and Jacala, Ba- rranca de San Vicente, near km 238, Moore & Wood 4435 (Gu), Moore & Wood 5065 (GH, MEXU); Real del Monte, Matuda 18883 (mexu), Zempoala, Benitez S36 (ENCB, MEXU). PUEBLA: Puebla- Veracruz border, Cumbres de Aculcingo, Boege 1953 (GH, MEXU). Veracruz: Puerto del Aire. Rosas 802 (GH, Mexu); El Puerto, Sharp 44703 (MEXU). a RRERO: Omiltepec, Halbinger s.n., November, 1975 (mexu). Cutaras: San Cristébal . Breedlove 11903 (mexu), Ghiesbreght 749 (Gu). Without definite locality,' ae 1088 1089 (Gu). — This widespread but rare, moderate-sized, scarlet-flowered Sa/via is one of the numerous attractive species in Mexico. Plants from Veracruz are glabres- cent, those of Puebla and Hidalgo very pubescent, and those of Guerrero intermediate. The number of flowers is variable: plants from a and Chiapas bear more than do those of Puebla, Veracruz, and Guerrero. Epling (1939) classified Salvia tubifera Cav. (including S. ee Willd.) in sect. TUBIFLORAE Epling, and SS. venosa and S. excelsa in sect. CURTIFLORAE ‘| do not know the definite localities for these two numbers. Fernald (1900) referred them to Zempoala in Hidalgo (Real del Monte is in Municipio Zempoala). Epling (1939), however, thought that one came from the Valley of Mexico, and the other from Real del Monte: he did not say which number was from where. 1984] RAMAMOORTHY, SALVIA 143 Epling. In general morphology S. tubifera resembles S. nervata Martens & Gal. of sect. CURTIFLORAE more than it does any members of sect. TUBIFLORAE. ACKNOWLEDGMENTS I thank Dr. P. F. Stevens for making available facilities in the Harvard University Herbaria, Dr. F. Chiang (Instituto de Biologia, Mexico) for the Latin diagnoses, Dr. J. Rzedowski (Escuela Nacional de Ciencias Biol6gicas, Mexico) for checking the Latin, Mr. D. Neill (Missouri Botanical Garden) for a partial review of the text, Dr. S. Castroviejo (Director, Instituto de Botanico A. J. Cavanilles, Madrid) for a gift of the photographs of types of Salvia spp. de- scribed by Cavanilles, and Ms Elvia Esparza (Instituto de Biologia, Mexico) for the illustrations. LITERATURE CITED BENTHAM, G. 1848. Labiatae. Jn; A. P. pt aa a 12: 27-603. CAvANILLES, A. 1791. Salvia tubifera. Ic. Des ee C. 1939. . revision of Salvia ae ee Repert, Spec. Nov. Regni g. Beih. 110: 1-383. ——_., oda, Se a notes on American Labiatae—I. Bull. Torrey Bot. Club 67: 504-534. ——.. 1951. Supplementary notes on American Labiatae—V. Brittonia 7: 129-142. ——. 1957. Supplementary notes on American Labiatae— VI. [bid. 8: 297-313. & = A AM. 1962. A new species of Sa/via from Mexico. Bot. Mus. Leafl. 20: 75, FERNALD, 7 s 1900. A synopsis of the oe and Central American species of Salvia. Proc. Amer. Acad. Arts 35: 489- Hemstey, W.B. 1882. Labiatae. /n: F. D. ee. SALVIN, eds., Biologia Centrali- Americana 2: 541-574 STANDLEY, P., & L. Wittiams. 1973. Labiatae. Fieldiana Bot. 24: 237-317. an INSTITUTO DE BIlOoLOGiA, DEPARTAMENTO DE BOTANICA APARTADO Postat 70-233, CluDAD UNIVERSITARIA DELEGACION COYOACAN 04510 Mexico, D.F., MEXICO BARRINGER, CUBITANTHUS 145 CUBITANTHUS, A NEW GENUS OF GESNERIACEAE FROM BRAZIL KERRY BARRINGER RUSSELIA ALATA Cham. & Schlecht. was originally assigned to the Scroph- ulariaceae and was included as such in the Flora Brasiliensis (Schmidt, 1862). Bentham (1846) noted that this species was not a Russelia L. f. but did not suggest an alternative placement until 1876, when he included it under 4ne- tanthus Hiern (Gesneriaceae) in Genera Plantarum. In all, Bentham included four species in Anetanthus; of these, Howard (1975) retained 4. gracilis Hiern and A. alatus (Cham. & Schlecht.) Durand & Jackson and transferred the others to different genera. Wiehler (1976) also reviewed the genus but found little relationship between 4. gracilis and A. alatus. He retained A. gracilis in a monotypic Anetanthus to which Skog (1982) has only recently added a second species. Wiehler rejected 4. a/atus as a gesneriad and suggested that it be placed somewhere in the Scrophulariaceae. Although Wiehler cited the type at Kew (Sellow s.n.), it appears that most of his observations are based on a plate published by Schmidt (1862) that misrepresents critical features of the species: the nectary is omitted, the placentation 1s shown as axile, and the seeds appear angular. Dr. Mattos Silva, of the Centro de Pesquisas do Cacau (CEPEC), has recently made material of this rarely collected species available to me. Study of these specimens confirms Bentham’s contention that the species should be included in the Gesneriaceae but shows that it is distinct from 4nefanthus and merits generic status. Cubitanthus Barringer, gen. nov. Herbae decumbentes pilosae, caules alati, radices fibrosae saepe adventitiae. Folia opposita petiolata serrata nervis pinnatis. Flores solitarii axillares ebrac- teolati, calyx quinquelobus lobis liberis lanceolatis acutis interne glabris, corolla quinqueloba bilabiata labio superio subintegro labio inferno trilobo ad basem villoso, stamina 4 geniculata, antheris cohaerentibus, discus integer, ovarium superum uniloculare placentatione parietali. Capsula septicida bivalvata, se- mina obovoidea striata. — Decumbent herb with long multicellular hairs; stem 4-winged; roots fibrous, often adventitious. Leaves opposite; petiole slightly winged; blade serrate, ve- nation pinnate. Flowers solitary in leaf axils; pedicel slightly winged at base, ebracteolate; calyx 5-lobed to base, the sepals lanceolate, acute, adaxially gla- © President and Fellows of Harvard College. 1984. Journal of the Arnold Arboretum 65: 145-147. January, 1984. 146 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 brous; corolla cylindrical, 5-lobed, bilabiate, the upper lip subentire, not re- flexed, the lower lip 3-lobed, densely villous at base; stamens 4, geniculate, adnate to base of corolla, the anthers bilocular, coherent in pairs; disc entire, annular; ovary superior, the placentation parietal, the style elongate, the stigma entire and slightly enlarged. Capsule septicidal, 2-valved, included within per- sistent calyx. Seeds ovoid, slightly striate. Type. Cubitanthus alatus (Cham. & Schlecht.) Barringer. EtymMo.oay. From the Greek kubitos, bearded, and anthos, flower, in reference to the densely villous lower lip of the corolla. Cubitanthus alatus (Cham. & Schlecht.) Barringer, comb. nov. _ alata Cham. & Schlecht. Linnaea 3: 3. 1828. Tyee: Bran, loc. ignot., Sellow Be MHONGE ype, | B iene ee: lectotype (here designated), k am. & Schlecht.) Bentham & J. D. Hooke: ex Durand & Jackson, 1893. Index Kew. I: ce SPECIMENS EXAMINED. Brazil. BAnIA: Itabuna, Ilheus, CEPEC, regiao de Mata Higrophila Sul Baiana, 15 Sept. 1981, Hage, Brito, & Argolo 1328 (CePrc, F); Itajuipe, San Antonio, 18 km do S de Itajuipe, 3 Feb. 1970, Dos Santos 554 (CePEc, F). Cubitanthus alatus is found as a weed in cacao plantations, but is known only from the cities of Ilheus and I[tajuipe in Brazil. It seems likely that it will be found more extensi vely in the coastal forest zone. The lectotype lacks locality data, as is typical of specimens collected by Sellow, but Urban (1906) reported that Sellow had collected in the coastal forests of Bahia in 1817 and it 1s likely that the specimen was collected at that time. The parietal placentation, disc, coherent anthers, and striate seeds are suf- ficient to place Cubitanthus in the Gesneriaceae. While these characters are also found in the Scrophulariaceae, they never occur in combination with each other. Both Anetanthus and Cubitanthus are decumbent herbs with entire discs and septicidal capsules. Anetanthus differs in having flowers in long-pedun- culate cymes, glandular-pilose calyxes that also have short glandular hairs on their inner surfaces, a bilobate upper corolla lip, stomatomorphic stigmas, and flattened, winged seeds. Cubitanthus is unique in having a winged stem and a densely bearded lower corolla lip. Both Cubitanthus and Anetanthus are best assigned to the Beslericae sensu Wiehler (1976). ACKNOWLEDGMENTS I would like to thank Mattos Silva for the generous gift of specimens. William Burger, Michael Huft, and Michael Nee helped with the earlier drafts. Laurence Skog kindly read and commented on the manuscript. REFERENCES BENTHAM, G. 1846. Scrophulariaceae. Jn: A. pe CANDOLLE, Prodr. Syst. Nat. Regni Veg. 10: 186-586. 1984] BARRINGER, CUBITANTHUS 147 ——.. 1876. Gesneriaceae. Jn’ G. BENTHAM & J. D. Hooker, Gen. Pl. 2: 990-1025. Howarpb, R. A. 1975. The genus Anetanthus. Jour. Arnold Arb. 56: 364-368. ScHMIpT, J. A. 1862. Scrophulariaceae. Jn: C. F. P. von Martius, Fl. Brasil. 8: 233- 330 SkoG, L. E. 1982. New Gesneriaceae from Peru and Ecuador. Selbyana 7: 94-99, Urnpan, I. 1906. a itineraque collectorum botanicorum, Jn: C. F. P. voN MARTIUS, Fl. Brasil. 1(1): 1-154. WIEHLER, H. 1976. re report on the classification of Achimenes, Eucodonia, Gloxinia, Goyazia, and Anetanthus. Selbyana 1: 374-404. DEPARTMENT OF BOTANY Fietb Museum oF NATURAL HISTORY CHICAGO, ILLINOIS 60605-2496 JOURNAL OF THE ARNOLD ARBORETUM INSTRUCTIONS FOR AUTHORS General policy The Journal of the Arnold Arboretum is primarily a staff journal, and staff papers have priority. 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STREET 1 SALES remouox OE ALERS ANDO CARRIER a VENOOR COUNTER SALES none none fa Seo nee 2 MAIL SUBSCRIPTION 712 689 ¢ 107 712 689 SAMPLES COMPLIMENTARY AND OTHER FREE COPIES 4 4 € rorat 716 693 F COPIES NOT OISTRIBUTEO ‘ Once use LEFT OVER, UNACCOUNTED SPOILED 84 107 NG none none G TOTAL (SumolF FI a? - 800 800 SIGNATURE AND TITLE OF EDITOR PUBLISHER BUSINESS nil ay ie) the statements made by MANAGER. OR OWNER ee. are correct and complete th tke ee Wha—opf PS Form IS26 July 1981 ( (Page!) Journal of the Arnold Arboretum January, 1984 CONTENTS OF VOLUME 65, NUMBER | Temple Clayton, Chemist and Amateur Botanist, 1914-1978. BRNICE Ur SCHUBERT 3044054444 edoecephaaedweiase meen eens I The Zingiberales (Cannaceae, Marantaceae, and Zingiberaceae) in the Southeastern United States. GEORGE K. ROGERS .. 0.0.00... 000000 ec ee 5 A Monograph of Diphylleia (Berberidaceae). TSUN-SHEN YING, SUSUMU TERABAYASHI, AND DAVID E. BOUFFORD | s434h54.04 cui sieaine op pnw ded 6b nadens 24545 eee kos 57 A Possible Magnolioid Floral Axis, Loishoglia bettencourtil, from the Upper Cretaceous of Central California. et se oy hbo og Ae wed oh he REA ROS 95 Systematics and Palynology of Picrodendron: Further Evidence for Relationship with the Oldfieldioideae (Euphorbiaceae). W. J. HAYDEN, W. T. GiLuis, D. E. STONE, C. R. BROOME, AND ts Vee 454 coe nde erate esa e eden eda oe bee ansadeed 105 The First Species of Stauranthera (Gesneriaceae) from New Guinea, with General Notes on the Genus. Bd RL. aad Bebe alk oe a Sa de EA 129 Notes on Sa/via (Labiatae) in Mexico, with Three New Species. Tes Ps RAMAMOORTHY 3. i0lth Ans node baa Eb bee oa oieee ae 135 Cubitanthus, a New Genus of Gesneriaceae from Brazil. KERRY BARRINGER: once deedch6 5 needa deke ee Seah ee be Mew N eet 145 Volume 64, Number 4, including pages 491-665, was issued October 20, 1983. JOURNAL oF tie ARNOLD ARBORETUM HARVARD UNIVERSITY VOLUME 65 NUMBER 2 US ISSN 0004-2625 Journal of the Arnold Arboretum Published quarterly in January, April, July, and October by the Arnold Arboretum, Harvard University. Subseripuion price $50.00 per year, plus $5.00 postage for addresses outside of U.S.A Subscriptions and remittances should be sent to Journal of the Arnold Arboretum, P. O. Box 368, Lawrence, Kansas 66044, U.S. A. Claims will not be accepted after six months from the date of issue. POSTMASTER: send address aa to Journal of the Arnold Arboretum, P.O. Box 368, Lawrence, Kansas 66044, Volumes 1-51. reprinted. and some back numbers of volumes 52-56 are available from the Kraus Reprint Corporation, Route 100, Millwood, New York 10546, J EDITORIAL COMMITTEE S. A. Spongberg. Editor E. B. Schmidt, Managing Editor P.S. Ashton K. S. Bawa P, F. Stevens C.E. Wood, Jr. Printed at Allen Press, Inc., Lawrence, Kansas COVER: The stylized design appearing on the Journal and the offprints was drawn by Karen Stoutsenberger Second-class postage paid at Boston, Massachusetts, and additional offices. JOURNAL OF THE ARNOLD ARBORETUM VOLUME 65 APRIL 1984 NUMBER 2 FRUITS AND SEEDS OF THE CUNONIACEAE WILLIAM C. DICKISON THE DICOTYLEDONOUS FAMILY Cunoniaceae consists of woody plants almost totally restricted to the Southern Hemisphere. It has long been of evolutionary interest as a putatively primitive and basal or near-basal group in the large rosalean complex from which a number of families have perhaps been derived (Dickison, 1975a). Depending upon taxonomic interpretation, it contains be- tween 19 and 27 genera. As pointed out by Dickison (1980b), members of the family are morpho- logically diverse, and considerable difference of opinion still exists regarding both generic circumscription and intrafamilial and interfamilial relationships. For example, an overall similarity in gross floral morphology has recently led Hoogland (1979) to reduce four Australasian genera to synonymy with the previously monotypic Chilean genus Cal/dcluvia D. Don and, at the same time, to establish the new segregate genus Acsmithia Hoogl. Within the family, Engler (1928), in his treatment for Die Natiirlichen Pflanzenfamilien, recognized 26 genera distributed among five tribes. These five tribes have been maintained by Schulze-Menz (1964) in the most recent edition of Engler’s Syllabus. Avail- able evidence indicates, however, that Engler’s treatment does not accurately reflect evolutionary groupings within the assemblage It is becoming clear, moreover, that parallel and reticulate evolution have often produced superficially similar morphologies among cunoniaceous genera. It is equally clear that the previously available morphological and anatomical data have often led to inaccurate systematic conclusions. During the past several years, I have been engaged in a thorough morphological and anatomical study of the Cunoniaceae with three main objectives: a better circumscription of the family and an eventual clarification of the interrelationships among genera of Cunoniaceae; an elucidation of the relationships of the Cunoniaceae to other families: and a better understanding of the overall patterns and trends of evo- lution within the family. Toward these goals, earlier research has dealt with aspects of floral (Dickison, 1975a), foliar (Dickison, 1975b), nodal (Dickison, 1980a), and wood anatomy (Dickison, 1980b). Although the resulting data © President and Fellows of Harvard College, Journal of the Arnold Arboretum 65: 149-190. et 1984. 150 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 have provided further insight into the relationships and relative degrees of advancement of various taxa, additional detailed and comprehensive study of other aspects will be required in order to obtain a reasonably accurate picture of intrafamilial relationships. This paper describes the anatomy of both fruits and seeds. The very scanty embryological information available on the Cunoniaceae has been summarized by Davis (1966). Aside from the incomplete accounts provided in numerous regional floras, little has been written concerning fruit and seed structure of the family (see Corner, 1976). More recent contributions dealing with cunoniaceous embryology and seed structure include papers by Govil and Saxena (1976) on Weinmannia fraxinea, Prakash and McAlister (1977) on Bauera capitata Ser., Krach (1977) on general seed morphology in the Cunoniaceae, and Kennedy and Prakash (1981) on Ca/licoma serratifolia. Outline drawings of seeds were provided by Bernardi (1964) in his revision of Weinmannia. MATERIALS AND METHODS Fruits and seeds of 20 genera and close to 100 species were examined in the present study. Specimens studied are presented in TABLE 1. Preserved fruits and seeds, in various stages of maturation, were embedded in paraffin and cut on a rotary microtome. Sections were subsequently stained with a combination of safranin and fast-green. Additional seeds were gathered from mature fruits on herbarium specimens housed at A, BRI, CANB, DUKE, GH, L, NCU, NOU, and z (herbarium acronyms according to Holmgren, Keuken, & Schofield, 1981). For scanning electron microscopy, dried seeds were affixed to stubs with double- stick tape, gold coated, and observed directly. The concept used of familial and generic limits follows Hoogland (1960, 1979, 1981). Most seed terminology follows the usage of Corner (1976). Slides are deposited in the Department of Biology, The University of North Carolina at Chapel Hill. OBSERVATIONS DEHISCENT-FRUITED GENERA Fruits. Twelve genera are characterized by dehiscent fruits: renee Ben- tham (1 species), Acsmithia (14), Bauera Banks ex H. C. Andrews (3), Cald- cluvia (11), Callicoma H. C. Andrews (1), Cunonia L. (ca. 20), Geissois Labill. (ca. 17), Lamanonia Vell. (6 to 8), Pancheria Brongn. & Gris (ca. 28), Spi- raeanthemum A. Gray (6), Vesselowskya Pampan. (1), and Weinmannia L. (ca. 150). Fruits and seeds of Bauera were the only ones not examined. In Acsmithia and Spiraeanthemum each carpel of the apocarpous gynoecia matures into a follicular fruit (FiGuRE 1), while all other genera produce two- (to five-)valved, sometimes horned capsules that undergo septicidal dehiscence (FiGures 2, 4-7). In contrast with the capsules of other genera, those of Cunonia dehisce by means of an acropetal separation of carpels from the persistent central column of tissue (FIGURE 3). Fruit-wall surfaces range between glabrous 1984] TABLE 1. DICKISON, CUNONIACEAE Specimens of Cunoniaceae examined. geal COUNTRY OF TAXON COLLECTOR ORIGIN VOUCHER Acrophyllum australe (A. Cunn.) Hoogl. Stauffer et al. 5700 Australia A Acs ria densiflora (Brongn. McKee 2 New Caledonia L & ris) Hoogl. New Caledonia NCU pw Caledoni NCU New Caledonia z A. elliptica (Pampan.) Hoogl. Veillon 2844 New Caledonia NCU A. integrifolia (Pulle) Hoogl. New Guinea L A. pedunculata (Schlechter) Hoogl. Veillon 1079 New Caledonia NOU A. pubescens (Pampan.) Hoogl. Schmid 431 New Caledonia NOU A. pulleana (Schlechter) Hoogl. Brass 12017 New Guinea A, L A. reticulata (Schlechter) Hoogl. Galore & Wood, NGF 41005 New Guinea A, L A. undulata (Vieill.) Hoogl. Vieillard 2078 New Caledonia b, Aistopetalum multiflorum Schlechter Hoogland & Craven 10826 New Guinea CANB Anodopetalum biglandulosum Webb _ 3337 Tasmania BRI (Hooker) Hooker £ Caldcluvia australiensis (Schlechter) Hoogland 8538 Australia A ra Hoogl] C. brassii (Perry) Hoogl. Br 30592 New Guinea L feat: 29616 New Guinea New Guinea CANB (on (Blume) Hoogl. New Guinea 1 17084 Yew Gu A Solomon Is. BRI Bougainville Is. BRI Cc. clemensiae (Perry) Hoogl. Van Balgooy 969 New Guinea L Cc. fulva (Schlechter) Hoogl. Van Royen, NGF 18266 New Guinea L C. nymanii (K. Schum.) Hoogl]. New Guinea L New Guinea L Ne i Z Jew Guinea L New Guinea A C. paniculata (Cav.) D. Don Dombey 713 Chile L Eyerdam & Chile GH C. paniculosa (F. Mueller) Hoogl Thorne et al. 25938 Australia L Hoogland 1115 Australia CANB C. papuana (Pulle) Hoogl. Kalkman 5316 New Guinea L rosifolia (A. Cunn.) Hoogl. Wilkes Exp. s New Zealand GH Walker s-.n New Zealanc GH C. rufa (Schlechter) Hoogl. Hoogland & Pullen 6081 New Guinea Hoogland & | New Guinea CANB icoma serratifolia BH. C. Andrews Hoogland 11692 Australia A, CANB 295 Australia BRI, NCU ¥ ustralia A Ceratopetalum succirubrum C. T. White Hartley 10,96 New Guinea A Croft & Marsh, LAE 71177 New Guinea BRL Codia nitida Schlechter Dackison § 221 New Caledonia NCU Jaffre 222: New Caledonia NOU C, obcordata Brongn. & Gris Guillaumin & Baumann- New Caledonia A Bodenheim 13117 Cunonia atrorubens Schlechter Blanchon 649 New Caledonia NOU FRUIT + + + + + + + t+ + 152 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 65 TABLE | (continued). COUNTRY OF TAXON COLLECTOR ORLCIN VOUCHER FRUIT SEED C. balansae Brongn. & Gris New Caledonia NCI + ei C. capensis L. Lam & Meeuse 4656 South Africa l + C. Llenormandii Pampan, McKee 7975 New Caledonia L + C. macrophylla Brongn. & Gris Bernardi 9507 New Caledonia or: Webster & Hildreth 14493 New Caledonia GH + C. pseudoverticillata Guillaumin McKee 4410 New Caledonia 1 + C. pterophylla Schlechter Dickison New Caledonia NCL + + C. pulchella Brongn. & Gris Dickison 287 New Caledonia NCU + + Dickison 218 New Caledonia NCL + + C. purpurea Brongn. & Gris Dickison 156 New Caledonia NCU + Dickison 144 New Caledonia NCU + C, schinziana Duniker McKee 23112 New Caledonia NOU + Geissois benthamiana F. Mueller Australia 1 as G. biagiana (F. Mueller) F. Mueller Dockrill 34 Australia I + G, hippecastaneifolia Guillaumin New Caledonia NCU af a G. pruinosa Brongn. & Gris Dickison 217 New Caledonia NCU + ot G. stipularis A. C. Sm. Fiji Herb, 31297 Fiji NCL + + New Guinea A + ae Lamanonia ternata Vell. Irwin et al. 8603 Brazil GH + L, tomentosa (Cambess.) 0. Kuntze Balansa 4752 Paraguay L af Pancheria confusa Guillaumin illon 66 New Caledonia NOU + P. elegans Brongn. & Gris New Caledonia NCU + + P. hirsuta Vieill. New Caledonia NCU au + P, reticulata Guillaumin Bernardi 12768 New Caledonia 4 + P. robusta Guillaumin Dickison 248 New Caledonia NCU + + P. sebertii Guillaumin Franc 2109 New Caledonia A + Platylophus crifoliatus (Thunb.) D. Don Bolus 621 S. Africa A + + Wiche 460/461 S. Africa L + Pseudoweinmannia lachnocarpa Clemens 43294 Australia A + + Mueller) Engler Pullea glabra Schlechter 31156 New Guinea A of + P. stutzeri (F. Mueller) Gibbs Irvine 1253 Australia BRI e Schizomeria ilicina (Ridley) Schlechter Brass 12150 New Guinea A + S. ovata D. Don Hoogland 11684 Australia CANB + + Dick ison & Jessup 297 Australia BRI, NCU + + whitei Mattf. Dickison 205 Australia NCU f tT Spiraeanthemum katakata Seem, Aw Fiji A + An ith 4409 Fiji L + Ss. Morrison 25.6.1896 New Hebrides A ef S. samoense A. Gray Bristol 2136 Samoa L + elowskya rubifolia (F. Mueller) Pampan. 11482 Australia ! + hodde 3210 Australia CANB + + D'Allei: Australia L + Weinmannia affinis A. Gray A. C. Smith 7698 Fiji L + W. aphanoneura Airy Shaw Ashton, BRUN 1044 Borneo L + W. bangii (Rusby) Pampan. Buchtien s.n. Bolivia L + 1984] DICKISON, CUNONIACEAE 153 TABLE | (continued). COUNTRY OF TAXON COLLECTOR ORIGIN VOUCHER FRUIT SEED W. blumei Planchon Dickison 215 New Guinea NCU ie + oorders 7647B Java a Schmutz 4588 Lesser Sunda Is. L + W. celebica Koord. Van Balgooy 3809 Celebes L it W. clemensiae Steenis Clemens 50877 Borneo L + W, decora Tul. Bernardi 11961 Madagascar L, Z@ or W. denhamii Seem. Bernardi 13258 New Hebrides L + Bernardi 12986 New Hebrides P + W. descombesiana Bernardi Celebes L qj W. dichotoma Brongn. & Gris New Caledonia 2 + W. fraxinea (D. Don) Miq. Cult., Java Ly + W. glabra L. Venezuela I + W. hutchinsonii Merr. Philippines L + W. luzoniensis Vidal Philippines BO + W. mauritiana D. Don! Réunion I a W. monticola Daniker New Caledonia NOU + W. negrosensis Elmer Ramos, Philippine Pl: Philippines L. + 1287 W Bernardi 9881 New Caledonia I + W. parviflora Forster Gillett 2215 Marquesas 1 l + W. pinnata L. Sousa & Rico B118 Mexico NCU + W. pullei Schlechter Hoogland & § New Guinea z + W. purpurea Perry Kajewski 1738 Solomon Is. L + W. racemosa L. f. Philipson et al. 349 New Zealand NCU + i on. Cult., E + + W. richii A. Gray A.C 13 Fiji L + W. rubrinervis Ettingsh. 7 D'Alleizette s.n. Madagascar l + W, serrata Brongn. & Gris Bernardi 12816 New Caledonia i + W. subsessiliflora Ruiz & Pavon Vasquez A130 Peru DUKE + W. sylvicola A, Cunn. Van Steenis 22314 New Zealand l + W. tannaensis Guillaumin Bernardi 12909 New Hebrides L;- 2 + W. tinctoria Sm. 4516 Réunion d + W. trichophora Perry New Guinea i + W. trichosperma Cav. 37 Chile 1 + W vescot. Drake Van Balgooy 1796 Tahiti al + Isyn. W. biviniana Tul. Syn. W rutenbergii Engler. and tomentose. The calyx is either persistent or deciduous in the fruit, with both of these character states sometimes present within a single genus (e.g., Weinmannia). All of the plants have mature pericarps that are differentiated into an outer parenchymatous region and an inner region of rigid, thick-walled, and highly lignified cells (FiGures 19, 20, 22). The endocarp tissue is thicker along the 154 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 dorsal wall and becomes thinner in ventral regions. In older fruits of Cunonia, the endocarp may become separated from the outer parenchymatous layers. The lignified valve margins, which represent the halves of the original septa, are illustrated in FiGures 2 and 5. Pericarp thickness ranges from a low of four to eight cells (Weinmannia) to a high of 30 to 40 cells (Geissois). Fruit vas- culature mirrors the pattern of gynoecial venation (Dickison, 1975a), although in the mature fruit veins become ensheathed by sclerenchyma. Epidermal cells of the outer surface of the pericarp are generally rectangular in transection, with a moderately thick cuticle on the outside. Weinmannia racemosa has a conspicuously striate external pattern. Mature fruits of Cunonia pulchella are covered with six or seven layers of periderm and have scattered lenticels. The parenchymatous central region of the pericarp is composed of isodiametric cells that may become moderately thick walled in older fruits. Numerous brachysclereids are present in the mesocarp. Scattered “‘mucilagi- nous idioblasts” with frothy contents occur in C. balansae, C. pterophylla, Geissols hippocastaneifolia, and Pancheria confusa. Druses are common in Cunonia balansae, C. capensis, C. pterophylla, and Pancheria robusta. In Ac- smithia densiflora, Caldcluvia rosifolia, Geissois pruinosa, Pancheria gatopen- sis Vieill. ex Guillaumin, and Vesselowskya rubifolia such crystals are confined to the boundary cells between outer parenchyma and endocarp or to the septum. In some species of Cunonia, zonation is indicated in the parenchymatous region of the pericarp by peripheral cells that stain more darkly. The inner, multiseriate sclerenchymatous region of the fruit wall, which becomes prominent following fertilization, is composed of two distinct zones of variable thickness in which the elongated cells are oriented in opposite planes: cells of the outer zone are arranged with their long axes parallel to the long axis of the fruit, while those of the inner zone are at right angles. DEVELOPMENT OF THE SEED COAT. Seed coats at early stages of development were examined for a number of capsular cunoniaceous taxa, including Cunonia balansae (FiGure 131) and Pancheria robusta (FiGuRE 134). Mature ovules are anatropous, bitegmic, and as far as is known, crassinucellate (Davis, 1966). At an early stage of seed-coat development, the outer integument is composed of a tanniferous outer epidermis and one or two subdermal layers consisting of smaller, nucleated, less densely staining cells. The outer epidermis of the inner integument gradually becomes transformed into a sclerotized exotegmen in the mature seed coat; during early stages of development, this layer is composed of rather small, nontanniferous cells with prominent nuclei (FiGuREs 131, 134). Although distinction between outer and inner integuments is typically lost in the mature seed coat, examination of developmental stages during seed-coat maturation shows that seeds of the Cunoniaceae are exotegmic (sensu Corner, 1976); that is, the outer epidermis of the inner integument differentiates into the mechanical layer. The middle two to four layers of the inner integument, some of which may arise by secondary periclinal divisions, consist of com- paratively enlarged, nontanniferous or tanniferous cells that are often tangen- tially flattened. Additional layers may develop along the seed edges. The inner epidermis is typically made up of densely cytoplasmic, tannin-accumulating 1984] DICKISON, CUNONIACEAE 155 cells that undergo little enlargement. In older seeds the inner epidermis always stains very darkly (FiGureE 133). MATURE SEEDS. The taxonomic literature indicates that seeds range in length from small (ca. 0.4 mm) to relatively large (ca. 6 mm). I have found no ex- ceptions to these dimensions. Outline varies from narrowly elliptic to elliptic or lanceolate. Angular seeds occur in Cunonia (Ficures 44, 5 1). Seed numbers range from one to many per locule, with ovule abortion common (FIGURES 130, 132). In cunoniaceous seeds the major features that show significant vari- ation and that may be combined in different ways are type and distribution of epidermal appendages, shape and wall characteristics of epidermal cells, and occurrence, type, and abundance of warts and striations. Seed coats are either glabrous or papillate (tuberculate), and appendages are usually present in the form of wings or hairs. A hilar scar is clearly visible in most taxa (FIGURE 91), and an elongate raised raphe is generally present. Seeds are supplied by a single undivided raphal bundle (Ficure 135). A straight embryo is always embedded in abundant starchy endosperm (Ficures 135, 136). Wings occur on seeds of all examined specimens of Acsmithia, Callicoma, Cunonia, Geissois, Lamanonia, Pancheria, Spiraeanthemum, and Vesselow- skya, as well as on those of some specimens assigned to the genus Caldcluvia. Wings are without vasculature and vary in size, shape, and location. Those of Geissois are long, flattened structures confined to an upper terminal position on the seed (FiGuRE 37). The same condition is found in some species of Pancheria, such as P. confusa (Ficures 31, 32), and in Acsmithia vitiense (A. Gray) Hoogl. Terminal wings attached both chalazally and micropylarly—and also sometimes laterally—occur in Acsmithia, Caldcluvia, Cunonia, Lama- nonia, Spiraeanthemum, Vesselowskya rubifolia, and some species of Pan- cheria (Ficures 33, 39, 46, 49, 52, 55, 57, 58, 65, 71, 73, 81). When both distal and basal wings are present, the two often differ in length and shape. In a number of species of Acsmithia, Cunonia, Lamanonia, and Pancheria, wings sometimes extend along one or both sides of the seed body as lateral append- age(s) (Ficures 35, 39, 49, 51, 55). Such lateral wings may extend up to the entire length of the seed body and appear always to form on the hilar side. Wing size and shape are also quite variable, ranging from rather long, broad structures with rounded apices, to triangular, to short, narrow outgrowths. Structurally mature wings consist primarily of two epidermal layers; between these one to three layers of unorganized or crushed cells may occur. Hairy seeds characterize the large and widely distributed genus Weinmannia and the New Zealand and Australian species of Caldcluvia—C. australiensis (Ficure 61), C. paniculosa (FiGure 67), and C. rosifolia (FiGURE 62). Three principal patterns of hair distribution are present: scattered, sparsely distrib- uted, not confined to the ends of seeds (FIGURES 61, 62, 67, 87, 91, 102, 105, 112); in tuft restricted to the ends of seeds (FIGURES 83, 85, 88, 97, 100, 103, 106, 109); and abundant over the entire seed surface (FiGuRES 92, 96). Inter- mediate conditions exist in which seeds are predominantly apically comate, with hairs also occasionally present on the sides (Figure 108). Great variation also occurs in hair length and abundance, with seeds ranging from sparsely 156 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 hirsute in W. tinctoria (FiGuRE 91), W. trichosperma (Ficure 112), and W. bangii, to very tomentose in such species as W. pullei (FiGuRE 92) and W. rubrinervis (FiGURE 96). In W. richii tufts of long hairs are present at both ends of the seed, and short, almost papillalike hairs are scattered over the sides (Ficures 94, 95). Seed hairs are simple, unicellular, sometimes striate, thin- walled structures without specialized ends such as hooks or barbs and are typically curved and intertwined in the mature seed. Surface features of seeds belonging to this group of genera are likewise quite variable. Species differ not only in the more readily visible aspects of epidermal cell shape, size, outline, and wall characteristics, but also in the less conspicuous secondary features of external wall ornamentation. External epidermal cell walls are smooth, striate, warty, or a combination of striate and warty. Narrow cuticular striations mostly parallel with the long axes of epidermal cells have been observed in Acsmithia elliptica, Acrophyllum australe (FiGureE 80), Cald- cluvia celebica (FiGure 60), Callicoma serratifolia (Figure 78), and many species of Weinmannia such as W. decora, W. dichotoma, W. pullei, W. tan- naensis, W. trichosperma, W. richii, and W. purpurea (FiGures 84, 93, 113, 120). More or less irregularly oriented striations occur in Cunonia macrophylla, Geissois biagiana, Caldcluvia fulva, and C. papuana (Ficures 43, 48, 70, 72). Striations vary considerably in size and abundance (compare Ficures 111 and 113). A warty external surface has been noted in Cunonia purpurea, Vesse- lowskya rubifolia, Weinmannia blumei, W. pinnata, W. subsessiliflora, W. negrosensis, W. hutchinsonii, W. descombesiana, and W. celebica (Ficures 45, 82, 104, 116, 117). Some species such as W. affinis, W. monticola, W. serrata, W. sylvicola, W. luzoniensis, W. fraxinea, W. vescoi, and W. denhamii (FiGuRES 89, 98, 99, 101, 111, 114, 119) have both warts and striae. Warts vary in size and prominence and range in outline from circular (most commonly) to angular to sometimes rectangular (in W. /uzoniensis, Ficure | A markedly papillate (tuberculate) condition, due to the unicellular protru- sions of epidermal cells, distinguishes Acrophyllum australe (Figure 79) and Callicoma serratifolia (FIGURE 76), as well as Caldcluvia brassii, C. paniculata (FiGurE 64), C. papuana (Ficure 69), C. nymanii (Figures 73, 74, 135), C. clemensiae (FiGure 66), and C. celebica (FiGures 58-60). Hoogland (1979) has drawn attention to the fact that among species of Ca/dcluvia, only the mature seed wall is papillate. Seeds of Acrophyllum are unique in being com- pletely covered by papillae (FiGure 79). Each papilla is generally broad and flattened at the base, smooth at the apex, and with a striate cuticle on the sides (Ficure 80). Although the papillae of Ca/dcluvia are morphologically similar, they are scattered and cover the seed coat incompletely (FiGuREs 59, 64, 66) As in Acrophyllum, the papillae are flattened at the base, rounded apically, and striate, differing only in that the striae extend over the apex (FiGuRE 60). Callicoma serratifolia also produces papillate or tuberculate seeds with striate papillae (FiGuRE 78). This observation is in agreement with that of Kennedy and Prakash (1981). Scanning electron micrographs show that species of Pancheria have a dis- tinctive seed surface in which enlarged epidermal cells of various outlines, with smooth, rounded outer walls, are arranged in mosaic patterns. These prominent 1984] DICKISON, CUNONIACEAE [57 epidermal cells may completely cover the surface of both the wing and the seed body, as in P. elegans (FiGuRES 35, 36), or they may be scattered over the seed surface along with less prominent cells of irregular outline, as in P. sebertii (Ficure 34). Similar enlarged cells occur in Cunonia schinziana (Ficure 50) and C. macrophylla (FiGure 47). The surface of C. purpurea seeds, however, is composed of irregularly polygonal cells with thin, slightly raised lateral walls. Cells of irregular outline with moderately thick to thin lateral walls comprise the seed coats of Lamanonia (Ficures 39, 40) and Geissois stipularis (FIGURES 37, 38). The Australian species of Geissois have diagnostic surface patterns: the epidermal cells of G. benthamiana seeds have raised outer walls covered with an unorganized arrangement of pits and irregular ridges and grooves (Figures 41, 42), while those of G. biagiana seeds have dome-shaped outer walls covered with prominent striations that are oriented at a flat to steep angle in relation to the inner walls (FiGuRe 43). Narrowly rectangular surface cells with moderately thick and slightly raised lateral walls extending parallel with the long axis of the seed are present in Vesselowskya rubifolia (FIGURE 82). Structurally complex epidermal cells distinguish seeds of Callicoma serratifolia. These cells are thick walled and have a combination of thick surface ridges and striae forming intricate patterns of the type seen in FiGureE 77. The seed-coat surface of Spiraeanthemum katakata and S. samoense gen- erally has smooth, thin-walled, rectangular cells that tend to be without con- spicuously raised anticlinal walls and are oriented with their long axes parallel to the long axis of the seed (FiGure 53). Acsmithia elliptica, A. densiflora, A. integrifolia, and A. pulleana have a microreticulate sculpturing of small, square or rectangular to elongate cells with moderately thick, raised lateral walls (FIGURES 54, 56). It is particularly interesting that in some collections of Acsmithia, the surface pattern on the seed body is structurally different from that on the wings. In the taxa examined, surface cells on the wings are irregularly oriented, often interwoven, and fibrous looking—distinctly different in appearance from those on the body (Ficures 54, 55). This feature has not been observed in other cunoniaceous genera. The genus Caldcluvia includes plants with diverse surface features. In ad- dition to the papillate condition described previously, the winged seeds of such species as C. brassii and C. paniculata have irregularly rectangular, hexagonal, or polygonal epidermal cells. Among the New Guinean species, C. nymanii appears distinctive in lacking obvious surface striations (FIGURE 75). Caldcluvia australiensis, C. rosifolia, and C. paniculosa—species with hairy seeds—have surface cells with raised and irregularly undulate anticlinal walls (FiGures 62, 63, 68, 69) Seed-surface patterns in Weinmannia are the most diverse in the family. In addition to the diversity discussed earlier, presence and distribution of stria- tions and warts, and epidermal cell shape and wall characteristics are also quite variable. A reticulate surface composed of irregularly polygonal cells with mod- erately thick and raised radial walls occurs in W. pinnata (FiGure 90) and W. subsessiliflora (FiGure 87). A more irregular pattern of rectangular cells, in places approaching a rugulate condition, occurs in W. decora (FiGure 120) and W. tinctoria (FiGurE 91). Vertically elongate cells of somewhat irregular outline 158 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 are present in a number of species. Variations in the appearance of this con- dition are illustrated in FiGures 86, 107, and 118. In FiGure 86 fine bars can be seen connecting the parallel ridges. A series of low, rounded ridges inter- spersed with small, spherical or polygonal warts is characteristic of W. mon- ticola (FiGuRE 89). The seed-coat surface of W. tannaensis consists of a series of undulating, usually longitudinally extended, low ridges and valleys (FIGURE 84); two ridges lying in close proximity are often connected by strands of wall material. The most common surface pattern in Weinmannia seeds is one of vertically elongated epidermal cells with raised, undulate anticlinal walls (Ficures 95, 97, 102, 106, 107, 114-117). Low, irregularly undulating ridges are present in W. /uzoniensis, W. fraxinea, and W. denhamii (Ficures 101, 110, 111, 119). These ridges vary in height and width. Histologically, the mature seed coat in these genera is composed of a cuticle and (four or) five (or more) layers of cells (FiGURES 23-26, 28). Four layers distinguish both Acrophyllum australe (FiGURE 30) and Callicoma serratifolia, whereas between six and eight cell layers characterize Geissois. This results in a three-layered testa and a one-, two-, or many-layered tegmen. Epidermal cells are cutinized, thin- or moderately thick-walled, and rectangular in tran- section. Surface cells in Geissois and Pancheria are typically enlarged and are often radially elongated as viewed in transection, with curved or dome-shaped outer walls (Ficures 41, 42). In fully mature seeds of Geissois, the enlarged epidermal cells covering the wings may become very thick walled and sclerotic. Similarly enlarged cells also occur on the surface of some Cunonia seeds. The subepidermal layer 1s composed of rectangular or compressed, thin-walled, nonlignified cells. Among species examined, Acrophyllum australe, Caldcluvia rosifolia, Cunonia pulchella, Geissois stipularis, Pancheria confusa, P. gato- pensis, and Vesselowskya rubifolia have a crystalliferous hypodermis (FIGURES 25, 28). Each crystal cell contains a solitary, angular crystal, although infrequent druses were observed in G. stipu/aris. Beneath the hypodermis a single layer of thick-walled, lignified, sclerotic cells is present in all taxa. These cells com- pose the only mechanical layer of the seed coat and are oriented longitudinally. Some variation exists among genera with regard to the shape and type of wall thickening of the sclerenchymatous cells as viewed in transection. While scle- renchyma cells are most commonly cuboidal or shortly radially elongate, with moderately thick walls, those of Vesselowskya rubifolia are small in diameter and have very thick walls (FiGuRE 28). Seeds of Acsmithia densiflora (FIGURES 23, 136), Caldcluvia rosifolia, and Weinmannia racemosa have a sclerotic layer composed of cells with thickened inner and radial walls (U-shaped thickenings). Cells that are circular in transectional outline and have very thick walls occur in Acrophyllum australe (FIGURE 30). The tegmen consists of thin-walled, rect- angular or compressed (crushed) cells, with the innermost layer always filled with a darkly staining substance. INDEHISCENT-FRUITED GENERA Nine genera are characterized by indehiscent fruits: Aistopetalum Schltr. (2 species), Anodopetalum A. Cunn. ex Endl. (1), Ceratopetalum J. E. Sm. (6), 1984] DICKISON, CUNONIACEAE 159 Codia J. R. Forster & J. G. A. Forster (ca. 10), Gillbeea F. Mueller (2), Platy- lophus D. Don (1), Pseudoweinmannia Engler (2), Pullea Schltr. (3), and Schi- zomeria D. Don (ca. 18). The fruits of these genera range from indehiscent capsules, drupes, and berries to the winged fruits of Gillbeea (FiGures 8-14). Seed structure is also quite variable. I have found it most convenient, therefore, to describe each genus or group of genera individually. The most common fruit type is an indehiscent, unilocular or bilocular cap- sule, characteristic of Ceratopetalum, Codia, Pseudoweinmannia, and Pullea. Fruits of Ceratopetalum, Codia, and Pullea may be derived from gynoecia with inferior or half-inferior ovaries, resulting in a fruit wall that is partially of hypanthial origin. Fruits of Ceratopetalum are bilocular, one- or two-seeded, and surrounded by enlarged, stellately spreading sepals (FiGure 11). Up to four seeds occur in Codia and Pseudoweinmannia fruits, but only a single fertile one may be present in those of Pullea glabra var. glabra. Mature fruit walls range from glabrous to densely tomentose, as in Codia (FiGure 14) and Pseu- doweinmannia, respectively (FIGURES 12, 13). Pericarps of Ceratopetalum and Pullea are differentiated into outer exocarp, middle mesocarp, and inner fibrous endocarp. Internal to the epidermis is a chiefly parenchymatous middle region that, in both genera, contains scattered brachysclereids. The sclereids in P. glabra are thin walled, whereas those in C. succirubrum are conspicuously thick walled and pitted. Enlarged “‘mucilage” cells are also present in Pullea. The generally narrow, fibrous, lignified endocarp is composed of thick-walled elements arranged in two zones, one oriented vertically and the other horizontally. Immediately external to the inner scler- enchymatous region in fruits of Ceratopetalum is a uniseriate prismatic layer in which each cell contains a solitary angular crystal. The pericarp of Pseudoweinmannia also contains a prominent crystalliferous layer outside the lignified, multiseriate inner zone composed mostly of tan- gentially elongated fibrous cells. Mature fruits of Pseudoweinmannia develop characteristic placental proliferations that, at maturity, may largely fill the locules between seeds (FiGuRE 21). Proliferations appear to be derived from the inner pericarp wall as well as from the placentae and have a folded ap- pearance in sectional view; the seeds are situated between the folds. The pla- centae are composed of thin-walled parenchyma cells, although the lateral peripheral regions consist of two or three layers of moderately thick-walled elements that resemble endocarp tissue. The outer surface of the placenta is composed of one or two layers of conspicuously enlarged, elongated cells that are oriented radially. The capsule wall of Codia nitida is composed of an outer layer of small, irregularly shaped epidermal cells and one to three layers of larger, moderately thick-walled hypodermal cells. Internal to the hypodermis is an interrupted ring of very thick-walled fibrous cells, some of which may surround the fruit vasculature. The inner half of the pericarp is constructed of loosely arranged aerenchyma, but a distinct sclerenchymatous endocarp 1s absent (FiGures 144, 145). Seeds of Codia and Pullea tend to be rather small (less than | mm long) and 160 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 are more or less narrowly ellipsoid to ellipsoid (FiGures 121, 123, 124). Pseu- doweinmannia produces slightly larger, spheroidal seeds with a somewhat lobed appearance (FiGure 125). Seeds of Codia and Pullea are essentially without wings, although rudimentary terminal wings can be recognized in some spec- imens. Arillike appendages characterize seeds of Pseudoweinmannia lachno- carpa. These appear as a ring of rather small, distinct, localized outgrowths from the exostomal region of the testa (FIGURE 126). In the absence of devel- opmental observations, the origin and function of these structures is uncertain, although the overall structure has the general appearance of an elaiosome. The surface of Pseudoweinmannia seeds is smooth, composed of nonorna- mented epidermal cells with a polygonal outline (FiGurE 125). Seeds of Codia nitida, C. obcordata, Pullea glabra var. glabra, and P. stutzeri have a reticulate external surface composed of cells with moderately raised anticlinal walls and a polygonal or, more commonly, reticulate outline (FiGuRE | 22). Fine striations can be observed on the surface of C. nitida. Seed coats of Codia and Pullea are thin and undifferentiated. Pigmented outer and inner epidermal layers bound a central region of one or two layers of thin-walled cells. Mechanical cells are absent (FiGuRE 29). Outer epidermal cells have irregular outlines and outer walls that are dome shaped in transection. Seeds of Codia are supplied by a tenuous, undivided vascular bundle extending to the chalazal region, while those of Pullea glabra have a single bundle that subdivides and terminates within the chalaza. In contrast with the seed coats of the two previous genera, those of Pseu- doweinmannia lachnocarpa are composed of multiple layers with a centrally positioned zone of mechanical cells. Very enlarged outer epidermal cells lack tannin and in the apical regions of the seed are often radially elongated to the surface; epidermal cells typically have thickened outer and radial walls. Beneath .the epidermis are two layers of thin-walled cells of irregular size and shape. Internal to this hypodermis is a uniseriate zone of small, relatively thin-walled, mechanical cells. The multiple tegmen is composed of approximately seven or eight layers, including the inner, tanniferous epidermis. Seeds are vascularized by a single bundle that terminates within the chalazal region and shows no evidence of branching. Seed coats of Ceratopetalum succirubrum (FIGURE 27) develop a thick-walled outer layer of pigmented cells that are rectangular in transectional outline and covered by a cuticle. Beneath the epidermis are two or three layers of pigmented cells, including a layer containing large, scattered, prismatic crystals. The crys- talliferous layer overlies a well-developed uniseriate region of sclerenchyma. The fibrous sclerenchymatous elements are of variable width and outline but are generally rather wide with very thick, lignified, pitted walls. The remaining tegmen contains five or six indistinct layers of compressed and pigmented cells. I was not able to interpret the pattern of seed vasculature from the material at hand. The genus Schizomeria is characterized by ellipsoid to ovoid, one-locular, one-seeded, drupaceous fruits (FiGures 8, 141). The pericarp is strongly dif- ferentiated into an outer exocarp, an extensive fleshy mesocarp, and an inner stony endocarp. The relatively thin exocarp is composed of a surface layer of 1984] DICKISON, CUNONIACEAE 161 small, rectangular, thick-walled epidermal cells and a single hypodermal layer consisting of more or less rectangular thick-walled cells. The fleshy mesocarp contains very thin-walled parenchyma interspersed with numerous isolated brachysclereids and sclereid nests. Sclereids are thick-walled, highly pitted, and occasionally crystalliferous with a solitary druse in the cell lumen. The endocarp is constructed of interwoven, tangentially elongated fibrous elements. Numer- ous small prismatic crystals are present throughout the inner mesocarp and endocarp. Major vascular bundles are confined to the fleshy mesocarp. The seed coat is tightly compressed against the inner pericarp wall, with the outer seed surface containing narrow grooves and ridges (FiGuRE 129). Beneath a cuticular layer, the testal epidermis is composed of darkly staining cells that are mostly square in transection. Cells in the two subepidermal layers are rectangular and thin walled. Beneath the bilayered hypodermis 1s a distinctive uniseriate zone of columnar mechanical cells with unevenly thickened walls resembling hourglass cells (FiGurE 142). Immediately internal to the scleren- chymatous layer are one or two layers of small crystalliferous cells. These cells merge with the multiple-layered tegmen mesophyll that is derived by periclinal division in the fertilized ovule and is composed of at least 20 layers of enlarged, very thin-walled cells. Inner epidermal cells are rectangular in outline and very tanniferous (FiGurE 141). A single large raphe bundle subdivides near the chalazal end of the seed into a system of veins that terminates at the chalaza. Fruits of Aistopetalum are four- to six-loculate, ovoid drupes. Each locule potentially contains a single oblong, pendulous seed (Ficures 137, 138). Mature fruits of A. viticoides Schltr. average 20 mm in length and 13 mm in diameter, with seeds about 6 mm long (Hoogland, 1960). Two immature fruits of Aistopetalum multiflorum were sectioned; these were 6 mm in diameter and 8 mm in length. The immature pericarp of the fruits 1s differentiated into exocarp, mesocarp, and endocarp on the basis of cell shape and differential retention of stain. The epidermis of the uneven outer surface of the pericarp, composed of small, rather irregularly shaped cells, is covered by a thin layer of cuticle. The hypodermis consists of five to seven layers of small cells with slightly thickened walls. The major portion of the pericarp contains isodiametric parenchyma cells of various sizes. Cells of the inner mesocarp are densely tanniferous. While immature, the endocarp is distin- guished by highly tanniferous tissue (FIGURE 139). As the fruit matures, seven to ten layers of tangentially elongated elements adjacent to the locules become relatively thick walled and differentiate into a very hard endocarp. Crystals are absent from the pericarp in my materia The immature seed coat is tightly compressed against the endocarp wall. The outer epidermis is densely pigmented and is composed of cells that are more or less square or rectangular in transectional outline. The remaining testa 1s generally made up of six to eight layers of tanniferous, isodiametric, thin-walled cells. A uniseriate layer composed of cells having very narrow diameters, an- gular outlines, and moderately thick walls separates testa and tegmen. The multiple-layered tegmen consists of six to eight layers of large, thin-walled cells that are often radially aligned, as well as an inner, tanniferous epidermis. Crystals are absent from the seed coat. A single vascular bundle enters the seed, 162 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 extends through the parenchymatous testa (FiGure 139), and branches freely within the chalazal region to form numerous veins that become arranged within the chalazal end of the seed (FiGureE 140). The third genus with fleshy fruits is Anodopetalum. The pericarp of Ano- dopetalum, in contrast with those of Schizomeria and Aistopetalum, is unlig- nified and not strongly differentiated. The fruit is therefore a berry. An outer exocarp 1s characterized by a layer of small, rectangular epidermal cells and up to six subepidermal layers of collenchyma. An extensive mesocarp contains parenchymatous cells and numerous scattered, highly pitted, relatively thin- walled brachysclereids. Vasculature is confined to the middle zone and, as in other fleshy fruit types of the Cunoniaceae, is not associated with ensheathing sclerenchyma. The inner pericarp region lacks sclerenchyma but is recognizable by a ring of crystalliferous cells containing acicular, clustered crystals that are accompanied by small, irregular, angular forms (FiGure 143). The acicular crystals are more or less needlelike but have blunt ends, so they are somewhat intermediate between raphides and styloids. In my reexpanded material the seed coat is not well preserved, but it can be determined to be parenchymatous and to consist of more than ten cell layers. Epidermal cells are irregular in outline and often have dome-shaped outer walls. A distinct layer of sclerenchyma is absent. As in the other cunoniaceous genera with fleshy fruit, seeds are vascularized by a single vein that divides repeatedly in the lower half of the seed before ending within the chalaza. The indehiscent trilocular fruit of Gi//beea has three prominent wings that extend the length of the central fruit column and are formed by the lateral expansion of the carpel wall (FiGures 10, 15). Hoogland (1960) described the carpels as each containing a single seed, although only one fertile locule existed in each of the two fruits sectioned for this study (FiGuRE 15). The wings have a prominent reticulate venation, which in mature fruits is surrounded by massive sclerenchymatous sheathing. The pericarp is composed of small epidermal cells, a parenchymatous mesocarp, and a narrow, two- or three-layered endocarp composed of fibrous elements, which is separated from the parenchymatous outer region by a crystalliferous layer containing angular crystals. Seeds of Gillbeea are oblong and about 5-8 mm long. The seed coat in the material examined is rather homogeneous in construction, with three or four layers of relatively thick-walled, pigmented, greatly compressed cells. I have not been able to detect a sclerenchymatous layer. The testal epidermis is con- spicuously papillate or tuberculate (FiGURE 16). As in other Cunoniaceae, the projections represent unicellular extensions of epidermal cells (FiGuRE 17). As observed from sectioned specimens, the tuberculae are broad at the base, with rounded apices and with surface striae extending parallel to the long axis of the projection. Presumably, some of the projections are glandular and secrete the dark, resinous substances that cover the seeds of species of this genus. | have not been able to detect whether the single vascular bundle present at midlength in the seed undergoes subsequent branching. The fruit of the monotypic South African genus Platylophus is derived from a bicarpellate gynoecium and, at maturity, is bilocular but most commonly 1984] DICKISON, CUNONIACEAE 163 one seeded. Its basal region is turgid, and the apical halfis distinctly compressed (FiGureE 9). The fruit has been described as an imperfectly dehiscent capsule (Harvey, 1894) that remains closed for an extended period. I have seen no dehisced fruits. In Platylophus the pericarp is covered by epidermal cells with decidedly thickened and pitted outer and anticlinal walls. A narrow mesocarp is con- structed of very thin-walled parenchyma cells, although its veins are surrounded by sclerenchymatous sheathing. A thin, two- or three-layered, fibrous, lignified endocarp is separated from the outer mesocarp by a region containing angular crystals (FiGuRE 18). Seeds are elliptic, less than | mm in length, and sometimes with vestigial wings (FIGURE 127). The seed surface is finely reticulate, with the individual cells generally square or rectangular in outline and having slightly raised lateral walls (FiGuRE 128). The minute, spherical warts that cover the external cell walls in Figure 128 are apparently artifacts, since they were not present in all specimens examined. The narrow seed coat is relatively undifferentiated and lacks a distinct scle- renchyma layer. Internal to the pigmented outer epidermis is a hypodermis containing scattered angular crystals and two layers of compressed cells that are rectangular in transectional outline. The internal epidermis is similar to the outer one, except that the cells are smaller. DISCUSSION GENERAL SUMMARY OF FRUIT AND SEED STRUCTURE Plants belonging to the Cunoniaceae are readily divisible into genera with dehiscent fruits (either follicles or bilocular capsules) and indehiscent ones (dry—capsular or winged; or fleshy—drupes or berries). Fruit wings of Cera- topetalum are derived from an expanded and hardened calyx, whereas the winglike outgrowths of Gillbeea fruits represent extensions of the gynoecial wall. Dehiscent capsules open septicidally from the apex downward or, less commonly, from the base upward. Fruits of Ceratopetalum, Codia, and Pullea are typically derived from inferior or half-inferior ovaries. Fruit walls range from glabrous to densely pubescent. Most species of Cunoniaceae produce fruits in which the pericarp becomes differentiated into an exocarp, a mesocarp, and a lignified, fibrous endocarp. The endocarp portion ranges in thickness from about 50 «m in fruits of some species of Weinmannia to just over 600 um in plants of Geissois, with a distinct woody endocarp entirely absent only in Codia and the Heshy- fruited genus Anodopetalum. Codia is unique in having tous inner pericarp tissue (Ficgures 144, 145). The drupaceous fruits of Aistopetalum and (particularly) Schizomeria contain an extensive fleshy mesocarp. Fruit walls range in thick- ness from rather thin (four to eight cells) to very thick (forty cells); they typically contain scattered prismatic or druse crystals, or crystals (rarely acicular ones— Anodopetalum) restricted to the inner mesocarp and endocarp. In certain species of Anodopetalum, Ceratopetalum, Cunonia, Pullea, and Schizomeria, brachy- sclereids are distributed throughout all three regions of the pericarp. 164 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 65 Dehiscent-fruited genera form one to numerous viable seeds per locule, ranging in length from small (0.4 mm) to rather large (6 mm), with the ovular integuments uniformly developing wings or hairs at maturity. Seeds are gen- erally apically comate, although less commonly hairs are distributed in other patterns. The only exception to the above generalization is the monotypic Australian genus Acrophyllum, which forms small, conspicuously papillate (tu- berculate) seeds. External seed surfaces are quite diverse in ornamentation. Variable features include shape and wall characteristics of the epidermal cells, and occurrence, type, and abundance of striations and warts. Seed coats are derived from both ovular integuments and are thin—one to five cells thick, except in Geissois, where they may be up to seven cells in thickness. Testae are relatively undifferentiated and generally contain an outer and an inner tanniferous layer. All taxa have a single fibrous layer that, as far as can be determined, is always derived from the outer epidermis of the inner integument. A distinct crystalliferous layer is present in the hypodermis in certain species. Genera with indehiscent fruits produce one to several seeds per fruit and a variety of seed morphologies. Seeds range in size from small (less than | mm in length) in Codia and Pullea to rather large (up to 5-6 mm in length) in Aistopetalum and Schizomeria. Although seeds of all genera are devoid of external wall appendages in the form of wings or hairs, those of Pseudowein- mannia are unique among the Cunoniaceae in having an arillike outgrowth with the general appearance of an elaiosome. Seed surfaces are smooth (Pseu- doweinmannia), reticulate (Codia, Pullea), or eee (Gillbeea). e seed coats are thin (three or four cells), and either undifferentiated and lacking a mechanical zone (Codia, Pullea) or multiple layered and with a well-differentiated fibrous layer (Schizomeria, Pseudoweinmannia). Multiple- layered seed coats arise by secondary division within the integumentary layers of fertilized ovules. In Anodopetalum multiple seed-coat layers appear to be associated with the absence of a mechanical zone, but this observation requires additional documentation. As in seeds from dehiscent-fruited taxa, one or more crystalliferous layers may be present in certain species. All cunoniaceous seeds are vascularized by a single bundle, which in some fleshy-fruited genera undergoes branching within the chalaza. All Cunoniaceae have straight embryos that at maturity are embedded in an abundant endo- sperm. — EVOLUTIONARY TRENDS IN FRuIT STRUCTURE AND IN SEED STRUCTURE AND DISPERSAL The diversity encountered among fruits and seeds of the Cunoniaceae clearly represents a range of different levels of evolutionary adv t and dispersal methods. A major trend of specialization occurs toward the indehiscent con- dition. A summary of fruit types and methods of seed dispersal in the Cuno- niaceae is presented in TAsBLe 2. Although the bilocular, ventrally dehiscent capsule is the most common fruit type in the family, all available evidence indicates that apocarpy—and the resultant follicular fruits, such as those found in Acsmithia and Spiraeanthemum—is the primitive condition in the family 1984] DICKISON, CUNONIACEAE 165 TABLE 2. Summary of fruit types and methods of seed dispersal in Cunoniaceae. GENUS. DISTRIBUTION HABIT FRUIT TYPE AGENT OF ADAPTATION FOR DISPERSAL DISPERSAL DEHISCENT FRUITS Acrophyllum New South Wales Subshrubs Capsule t Seeds papillate Acsmithia Fiji, New Caledonia, Small to medium Follicle Wind Seeds winged Queensland, New Guinea, trees Moluccas Bauera E Australia Small shrubs, Capsule 2 ? subshrubs Caldeluvia Chile, New Zealand, Small to large Capsule Wind Seeds winged, Australia, Philippines, trees papillate, Celebes, Moluccas, hairy Guinea, Solomon Is. Bougainville Is llicoma E Australia Small trees, Capsule Wind Seeds winged, shrubs papillate Cunonia New Caledonia, Small to medium Capsule Wind Seeds winged South Africa trees, shrubs Geissois Fiji, New Caledonia Small to medium Capsule Wind Seeds winged trees 1onia Brazil, Paraguay Trees, shrubs Capsule Wind Seeds winged Pancheria New Caledonia Small t Capsule Wind Seeds winged hrubs Spiraeanthemum Samoa, Fiji, New Hebrides, Shrubs, small to Follicle Wind Seeds winged Solomon Is., New Britain, medium trees Bougainville Vesselowskya E Australia Small trees, shrubs Capsule Wind Seeds winged Small to medium Capsule Wind Seeds hairy Weinmannia INDEHISC Aistopetalum Anodopetalum Ceratopetalum Codia Gillbeea Platylophus Pseudoweinmann Pullea Schizomeria [ FRUITS S a Old and New World tropics New Guinea Tasmania E Australia, New Guinea New Caledonia Queensland, New Guinea South Africa Queensland, New South Wales Fiji, Queensland, Moluccas, New Guinea E Australia,.New Guinea, Solomon w trees, shrubs Tall trees Shrubs Large shrubs, small to large trees Shrubs, trees smal] Medium to large trees Medium to large Large ree trees Small to medium Drupe Capsule Capsule Wingec Capsule Capsule Capsule Drupe Animals(?) Animals(?) Wind Wind Water Wind (and a4 ants?) Animals(?) Fruits fleshy Fruits fleshy Fruits winged Fruits hairy Fruits winged Fruits turgid Fruits hairy, seeds with elaiosomes Fruits fleshy 166 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 and is not derived as suggested by Cuatrecasas (1970). The initial evolutionary step, therefore, involved the conversion ofa cluster of follicles into a septicidally dehiscent capsule. In all the genera considered to be primitive on the basis of floral morphology and wood anatomy, the seed 1s thin walled, has a single, lignified, fibrous layer, and acts as the sole dispersal unit. With the notable exception of Acrophyllum, all dehiscent-fruited genera produce seeds with structural modifications — either membranous wings or hairs—for dispersal by wind. Appendages have appar- ently been lost from the seeds of Acrophyllum, although the surface cells have prominent papillae or tuberculate outgrowths. In contrast with other taxa, Acrophyllum is a small (usually under | m tall) shrub or subshrub that has a very restricted distribution in the Blue Mountains of New South Wales, where it grows only on moist ledges in the immediate vicinity of waterfalls. Subsequent evolutionary advancement has resulted in major shifts in dis- persal methods. In the more advanced cunoniaceous taxa, the entire fruit has become modified for dispersal and protection, and there have been reductions in the number of seeds per fruit. Distinct trends of structural specialization in the fruit have resulted in densely pubescent (Codia, Pseudoweinmannia) and winged (Ceratopetalum, Gillbeea) fruit types that are associated with anemo- chory. Moreover, the two winged types of fruit represent different evolutionar trends since the wings are derived from different sources (in Ceratopetalum they derive from an enlarged and hardened calyx, whereas in Gillbeea they are gynoecial in origin). In Anodopetalum, Codia, and Bauera a lignified endocarp in the fruit wall has been lost (Prakash & McAlister, 1977). The adaptive significance of pericarp aerenchyma in Codia is unclear. Pseudoweinmannia, a tall rainforest tree from Queensland and New South Wales, is interesting in that its densely hairy, indehiscent fruits are undoubtedly scattered by wind, but its elaiosomelike seed appendages suggest that a secondary agent (perhaps an ant) may also be involved in dispersal. Trends have also occurred toward formation of drupaceous and berrylike fruits. Although method(s) of dispersal for these fruit types is unknown for the Cunoniaceae, such adaptations generally promote seed dispersal by animals. For Platylophus, which usually grows near streams and rivers, Ridley (1930) repeated earlier observations that the fruits are dispersed by flowing water. Ben-Erik van Wyk, of the University of Stellenbosch, has observed (pers. comm.) that the turgid capsules break off, leaving the entire pedicel on the inflorescence, and that capsules are frequently seen floating on water. The inflated fruit wall and the absence of dispersal structures on the small, thin- walled seeds may be reflections of this adaptation. Within the family there have thus been major adaptive shifts in dispersal agents— from wind to animal, and even water. Seeds have undergone concomitant changes. Accompanying the general loss of dispersal appendages, at least two distinct trends are evident in seed-coat structure: reduction in seed-coat thickness, including the loss of a mechanical layer (e.g., Codia, Platylophus, Pullea); and amplification of the seed coat by secondary division of integumentary cells in the fertilized ovule to form a 1984] DICKISON, CUNONIACEAE 167 multiple-layered seed coat (e.g., Aistopetalum, Schizomeria). A secondary in- crease in seed-coat layers may or may not be accompanied by loss of a fibrous layer. I cannot accept Krach’s (1977) suggestion that all seed coats in the Cunoniaceae were derived via a general trend toward reduction from a mul- tilayered ancestral condition. The various genera of Cunoniaceae are, therefore, good examples of what Corner (1958) and later Stebbins (1970, 1974) referred to as the “transference of function” with respect to the protection and dispersal of seeds. I am uncertain of the significance of the resinous coating on seeds of Gillbeea. It is of interest to note the general correlation between the production of indehiscent fruits with more specialized seed types and a more advanced wood anatomy. The genera Aistopetalum, Anodopetalum, Bauera, Ceratopetalum, Codia, Pseudoweinmannia, and Schizomeria all have vessel elements with exclusively or predominantly simple perforation plates (Dickison, 1980b). The only exceptions to this correlation are Gillbeea and Pullea, which have vessel elements with scalariform perforations. FRUIT AND SEED MORPHOLOGY AND INTRAFAMILIAL SYSTEMATICS I have previously pointed out (Dickison, 1980b) that evidence is accumu- lating to indicate that many of the tribes of Cunoniaceae, as defined by Engler (1928), are very unnatural assemblages that should be either reconstructed or abandoned. The morphology of fruits and seeds supports this opinion. Although the family can be divided into groups and subgroups, representing different levels of specialization, by the use of various characters, it is clear that fruit morphology should be important in any future subdivision of the family. Engler’s tribe Belangereae, containing the southwestern Pacific genus Geissois and the Neotropical Lamanonia, is from all indications a natural grouping. The two genera both lack petals and have numerous stamens and a bicarpellate superior gynoecium that matures into a septicidally dehiscent capsule contain- ing comparatively large winged seeds. Both also have a structurally advanced wood anatomy (Dickison, 1980b) that is correlated with other indices of ad- vancement such as fourth-rank leaves (sensu Hickey, 1971) and, at least in Geissois, multilacunar nodal anatomy (Dickison, 1980a). I emphasized in an earlier paper (Dickison, 1980b) that the tribe Spiraean- themeae is a very heterogeneous aggregation that includes the primitive genus Spiraeanthemum (including the segregate genus Acsmithia), with primitive wood anatomy, as well as the rather advanced genus Aistopetalum. Extreme diversity is also apparent in reproductive morphology. Spiraeanthemum (and Acsmithia) have follicular fruits with winged seeds, whereas the drupaceous fruit of Aistopetalum and the winged fruit of Gillbeea are both significantly more advanced and represent quite different dispersal methods. Although all species in this tribe have flowers with a mostly three- to five-carpellate gynoe- cium and a stamen number equal to or double the number of sepals, they differ with respect to such other characters as sepal aestivation (imbricate or valvate) 168 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 and gynoecium type (apocarpous to syncarpous). The genus Aphanopetalum appears to be best removed from the family for reasons previously outlined (Dickison, 1980b). Engler defined his Cunonieae, the largest tribe of Cunoniaceae, by the fol- lowing characters: stamens equal to or double the number of sepals; gynoecium superior, bicarpellate, with the carpels more or less fused ventrally in the region of the ovary; petals present or absent; sepals imbricate or valvate; and flowers not produced in a dense spherical head. On the basis of vegetative anatomy, this is the most diverse group, with genera representing quite different levels of evolutionary advancement (Dick- ison, 1980b). This is also true for the fruits and seeds of the tribe, which represent major differences in dispersal methods. Both dehiscent and indehis- cent fruits are present. The most frequent type is the dehiscent capsule, such as occurs in Acrophyllum, Caldcluvia, Cunonia, Vesselowskya, and Weinman- nia. There are also more advanced, indehiscent capsules (Ceratopetalum, Pseu- doweinmannia), drupes (Schizomeria), and berries (Anodopetalum). oogland (1979) has reduced the Australasian genera Ackama A. Cunn., Betchea Schltr. (including Sto//aea Schltr.), Opocunonia Schlitr., and Spiraeop- sis Mig. to synonymy with the monotypic Chilean genus Ca/dcluvia. The di- versity in wood structure among these taxa has already been discussed (Dick- ison, 1980b). Hoogland (1979) noted that the flowers and fruits of these species are too much alike to merit recognition as separate genera. The seeds, however, are different, as was pointed out by Hoogland. The three Australian and New Zealand species formerly included in Ackama all have hairy seeds and similar surface patterns, with epidermal cells having conspicuously undulate anticlinal walls. All other taxa in this complex have winged, frequently papillate seeds with striate or nonstriate (C. nymanil) epidermal cells. Hoogland (1979) further stated that Ca/dcluvia is similar to Weinmannia in flower and fruit, being most easily separated on the basis of inflorescence structure. In addition, relationship between Caldcluvia and Weinmannia is indicated by the occurrence of apically comate seeds in species of both genera. It should also be reemphasized that the only apparent characters that would argue against a merger of Weinmannia and Cunonia are those associated with fruit dehiscence and seed morphology. Seeds of Weinmannia are always hirsute, whereas those of Cunonia are uniformly winged. Since both character states currently exist in Caldcluvia, however, the strength of this distinction is weak- ened. In Engler’s scheme the tribe Pancherieae, which contains the three genera Callicoma, Codia, and Pancheria, is held together by flowers that are char- acteristically produced in tightly compacted, globose clusters. Xylem anatomy is quite variable among these genera, which also show both imbricate and valvate sepal aestivation, superior and inferior gynoecia, and dehiscent (Cal- licoma, Pancheria) and indehiscent (Codia) fruits producing seeds of quite different structure. The fruits of Codia, in contrast to those of Callicoma and Pancheria, lack a lignified endocarp, and its seeds also lack a mechanical layer. I think that a similar inflorescence type has evolved among diverse elements. The last tribe, Pulleae, contains the single genus Pullea, characterized by a 1984] DICKISON, CUNONIACEAE 169 rather primitive wood structure, apetalous flowers, imbricate sepals, and a bicarpellate, ‘‘half-inferior” ovary that matures into an indehiscent capsule. Seeds of Pullea, Codia, and Platylophus are similar in size, in lack of a fibrous layer in the seed coat, and in surface patterns. The systematic position of the Australian and Tasmanian genus Bauera Banks has been somewhat unsettled (Dickison, 1975c). Despite my earlier opinion to the contrary (Dickison, 1975c), recent evidence strongly favors a position in the Cunoniaceae (Bensel & Palser, 1975; Prakash & McAlister, 1977). Aspects of vegetative anatomy would place the genus among the more advanced genera within the family (Dickison, 1980b). The fruits of Bauera are dehiscent capsules with seeds that develop a multiple-layered tegmen and ap- parently lack a mechanical layer (Prakash & McAlister, 1977). This is an uncommon combination of features in the Cunoniaceae, occurring elsewhere only in Anodopetalum. A secondary increase in seed-coat layers and an absence of a thick-walled lignified cell layer are both, in my opinion, advanced char- acteristics among cunoniaceous seeds The use of scanning electron microscopy has revealed considerable variation in shape, pattern of hair distribution, and surface characteristics of the seeds. This is particularly true in the large and widely distributed genus Weinmannia. Although in a few cases features appear to be diagnostic for species, a much greater sample would need to be examined to assess fully the taxonomic use- fulness of seed morphology at the subgeneric and specific levels. COMMENTS ON THE RELATIONSHIPS OF CUNONIACEAE TO OTHER FAMILIES As reviewed by Dickison (1975a, 1980b), the Cunoniaceae have tradition- ally been placed in a basal position within the large order Rosales, in or near the saxifragaceous complex (Engler, 1928; Schulze-Menz, 1964: Cronquist, 1981). Families that have long been closely allied with the Cunoniaceae include the Saxifragaceae, Eucryphiaceae, Brunelliaceae, and Staphyleaceae. Since the seed structures of this complex are incompletely known, it would be premature to attempt a discussion of relationships based upon this criterion. However, a few observations can be made. The present study further substantiates the significant differences in seed- coat structure between the Cunoniaceae (including Bauera) and the Saxifra- gaceae that have been summarized by Prakash and McAlister (1977). Seed coats of the Cunoniaceae are uniformly derived from both integuments, where- as those of the Saxifragaceae apparently mature only from the outer integument, with the inner tegmen degenerating. Corner (1976) has indicated that there is a significant evolutionary distinction between families with exotestal seed con- struction (Saxifragaceae) and those with exotegmic seed construction (Cuno- niaceae). The additional important distinctions (relating to aspects of embryol- ogy) between these families require documentation. The small neotropical family Brunelliaceae, characterized by apocarpous, apetalous, diplostemonous flowers, in this respect resembles the cunoniaceous genera Acsmithia and Spiraeanthemum. Each carpel matures into a follicle in 170 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 which the pericarp becomes strongly differentiated into outer parenchymatous exocarp and inner woody endocarp (Eyde, 1970). The asymmetric growth of the ovary that displaces the style to the abaxial side—and the resulting spe- cialized fruit—is constant in all members of the Brunelliaceae and is not matched in any possibly related family (Cuatrecasas, 1970). Seeds of Brunellia lack dispersal appendages, and the testa is thick. Wood anatomy of the genus is at a higher evolutionary level than that of the more primitive genera of Cuno- niaceae (Dickison, 1980b). Over the years various individuals (Hallier, 1908; Linden, 1960: Whitmore, 1972; Thorne, 1976) have strongly emphasized tee between the Staphy- leaceae and the Cunoniaceae. Similarity is evidenced in the fibrous exotegmic seeds of the staphyleaceous genera Huertea Ruiz & Pavon and Tapiscia D. Oliver (Corner, 1976) and the exotegmic seeds of the Cunoniaceae. ACKNOWLEDGMENTS This study was supported by National Science Foundation grant DEB- 8025976. I wish to thank the following individuals who assisted me in various ways while I was collecting plants used in the present research: A. Sundaresan (Suva), P. Morat (Nouméa), G. McPherson (Nouméa), L. A. S. Johnson (Syd- ney), R. Coveny (Sydney), L. Jessup (Brisbane), B. P. M. Hyland (Atherton), and M. Galore (Lae). Thanks are also extended to the directors of the Harvard University Herbaria, the Queensland Herbarium, the Herbarium of the Bo- tanischer Garten und Institut fiir Systematische Botanik der Universitit Ziirich, and the Herbarium of O.R.S.T.O.M., Nouméa, for allowing me access to their collections. Dr. R. D. Hoogland kindly sent a large number of representative seeds from specimens housed at the Rijksherbarium, Leiden, and also offered valuable comments on the manuscript. Ben-Erik van Wyk (Stellenbosch) was very helpful in supplying information on the fruit of Platvlophus. Special thanks are also given to my colleagues P. G. Gensel, J. M. Massey, and A. E. Radford for kindly reading the manuscript and offering useful suggestions for improve- ment. The assistance of Cyndi Jones in slide preparation, Doug Wilson in obtaining SEM photographs, and Susan Whitfield in artistic preparation of plates is gratefully acknowledged. LITERATURE CITED eae C.R., & B. F. Parser. 1975. Floral eae in the Saxifragaceae sensu lato. V. Baueroideae and conclusions. Amer. J. Bot. 62: 688-694. ae L. 1964. Revisio generis aan Pars III: sectiones I1I-IV—V-VI (veteris orbis). Bot. Jahrb. Syst. 83: 126-221. Corner, E. J. H. 1958. Transference of function. J. Linn. Soc., Bot. 56: 3 ——. 1976. The seeds of dicotyledons. Vol. 1. lx + 311 pp. Cambridge a Press, Cambridge, Englan Cronguist, A. 1981. “An integrated system of classification of flowering plants. xvul + 1262 pp. Columbia Univ. Press, New CUATRECASAS, J. 1970. Brunelliaceae. FI. Neotrop. 2: 5- 189, Davis, G. L. 1966. Systematic embryology of the angiosperms. viii + 528 pp. John Wiley & Sons, New York. 1984] DICKISON, CUNONIACEAE Ll Dickison, W. C. 1975a. Studies on the floral anatomy of the Cunoniaceae. Amer. J. Bot. 62: 433-447. 19 Leaf anatomy of Cunoniaceae. J. Linn. Soc., Bot. 71: 275- 294, 1975c. Floral morphology and anatomy of Bauera. Phytomorphology 25: 69-75. 1980a. Diverse nodal anatomy of the Cunoniaceae. Amer. J. Bot. 67: 975- ~~ wn 981. Ob. Comparative wood anatomy and evolution of the Cunoniaceae. Aller- tonia 2: 281-321. ENGLER, A. 1928. Cunoniaceae. Jn: A. ENGLER & K. PRANTL, eds., Nat. Pflanzenfam. ed. 2. 18a: 229-262. Eype, R. H. 1970. Anatomy. Jn: J. CUATRECASAS, ed., Brunelliaceae. Fl. Neotrop. 2: Govit, C. M., & N. P. SAXENA. 1976. Anatomy and embryology of Weinmannia fraxinea Sm. (Cunoniaceae). J. Indian Bot. Soc. 55: 219-22 HALuerR, H. 1908. Uber Juliania, eine Terebinthaceen- Gattung mit ey und die wahren Stammeltern der Kdtzchenbliitler. Beih. Bot. Centralbl. 23: 81- Harvey, W.H. 1894. Saxifragaceae. Pp. 305-309 ee H. Harvey & O. SonDER: eds., Flora Capensis. Vol. 2. L. Reeve & Co., ent. Hickey, L. J. 1971. eae significance paren ey features in the woody dicots. (Abstr.) Amer. J. 58: 469. HoLMGREN, P. K., W. KEUKEN, oe E. K. SCHOFIELD. 1981. Index herbariorum. ed. 7. HooGLanp, R. D. 1960. Studies in the Cunoniaceae. I. The genera Ceratopetalum, wa Aistopetalum, and Calycomis. Austral. J. Bot. 8: 318-341. 979. Studies in the Cunoniaceae. II. The genera Caldcluvia, Pullea, Acsmithia, and Spiracantemn Blumea 25: 481-505. 19 s in the Cunoniaceae. III. Additional notes on Ceratopetalum and Acrophyllum. eae 213-216. Kennepy, M. J., & N. PRAKASH. 1981. A morphological and SL ey study of Callicoma serratifolia Andr. (Cunoniaceae). Austral. J. Bot. 29: 721-7 Kracu, J. E. 1977. Seed characters in and affinities among the ee a: K. Kusirzki, ed., Flowering plants. Evolution and classification of higher categories. Plant Syst. Evol., Suppl. I: 141-153. LinpeN, B. L. VAN DER. 1960. Staphyleaceae. Jn: C. G. G. J. VAN STEENIS, ed., FI. S SE PRAKASH, N., & E. J. McAuisterR. 1977. An embryological study of Bauera capitata with comments on the systematic position of Bauera. Austral. J. Bot. 25: 615-622 Riptey. H. N. 1930. The dispersal of plants throughout the world. xx + 744 pp. L. Reeve & Co., Ashford, Kent. ScCHULZE-MENz, G. K. 1964. Cunoniaceae. /n: H. Metcuior, ed., Syllabus Pflanzenfam. ds12: Steppins. G. L. 1970. Transference of function as a factor in the evolution of seeds and their accessory structure. Israel J. Bot. 19: 59- 9 Flowering plants: evolution above the species level. xviii + 399 pp. Be Iknap—Harvard Press, Cambridge, Massachusett THorne_, R. F. 1976. A Saas classification of the Angiospermae. /n: M. K. Hecut. W. C. STEERE, & B. WALLACE, eds., Evol. Biol. 9: 35-1 Wuitmore, T. C. 1972. Staphyleaceae. Pp. 446-448 in T. C. WHITMORE, ed., Tree flora of Malaya. Vol. 1. Longman Group Ltd., London. DEPARTMENT OF BIOLOGY Tue UNIversiry OF NORTH CAROLINA CHapeL HILL, NortTH CAROLINA 27514 T72 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 EXPLANATION OF PLATES PLATE I Ficures 1-14. Fruit morphology of Cunoniaceae. |, Spiraeanthe mum mac gillivrayi (Morrison 25.6.1896), x 5.5. 2, Weinmannia blumei (Dickison 215), x 5. 3, Cunonia macrophylla (Wi Biter & Hildreth 14493), x 1.5. 4,5, Pancheria sebertii ee 2109): mature fruit head, x 1.5; 5, individual f ven valve, x 3. 6, aes faa ie (Fiji Herb. 31297), x 1. 7, Acrophyltum australe (Stauffer et al. 5700), x . Schizomeria ilicina (Brass 12150), x — oe Ne ae (Bolus 621), x 3. see pa- puana (Brass 31815), x = . Ceratopetalum succirubrum (Hartley 10967), x 1.5.12 13, Pseudoweinmannia lac emi (C ee 43294), x 2.5: 12, mature fruit: 13, ma- ture fruit with tomentum partially removed. 14, Codia nitida (Dickison 221), x 2.5. PLATE II Figures 15-22. Fruit anatomy of Cunoniaceae. 15-17, Gillbeea papuana (Brass 31815): 15, transverse section of mature fruit; 16, transverse section of mature seed: 17. seed epidermis showing papillae. 18, Platylophus trifoliatus (Bolus 621), transverse sec- tion of pericarp. 19, Acsmithia de be (Dickison 220), transverse section of follicle. 0, mosa (cult., E), transverse section of capsule. 21, Pseudowein- mannia lachnoc a (Dickison & oe 294), transverse section of fruit showing pla- cental proliferations. 22, Gerssois stipularis (Fiji Herb. 31297), transverse section of capsule (only eae veins illustrated). PLATE III Ficures 23-30. Seed-coat structure in Cunoniaceae: 23, Acsmithia densiflora (Dick- ison 220), 24, Caldcluvia nymanii (Hoogland & Craven 11069), 25, Cunonia pulchella (Dickison 218), 26, Pancheria hirsuta (Dickison 227); 27, Ceratopetalum succirubrum (Croft & Marsh, LAE 71177), 28, Vesselowskya rubifolia (Schodde 3210), 29, Codia nitida (Dickison 221); 30, Acrophyllum australe (Stauffer et al. 5700) (CR, crystalliferous layer, CU, cuticle; EN, endosperm; EP, epidermis; IT, inner tegmen; S, sclerenchymatous layer; stippled cells pigmented). PLATE IV Figures 31-39. Scanning electron micrographs of seeds of Cunoniaceae. 31, Pancheria confusa (Veillon 66): 31, side view, x 9; 32, hilar region, x 20. 33, 34, sebertii (Franc 2109): 33, side view, x 9: 34 A Sure. x 40, 35, 36, P. elegans (Dickison 158): 35, oblique side view, x 25: 36 6, surface, x 50.37, 38, Geissois stipularis (Fiji Herb. 31297): 37, side view, x 8; 38, surface: x 22.39, Lamanonia ternata (Irwin et al. 8603), side view, x 6.5 PLATE V Ficures 40-48. Scanning electron micrographs of seeds of Cunoniaceae. 40, La- mManonia pineal orale 4752), surface, x oe 41, 42, Geissois Pie ey oa Guard, Lismore 3.1909), surface: 41, x 50; 42, x 500. 43. G. ae a (Dockrill oe surface, x 500. ee Cunonia purpurea (Dickison 144): 44, side view, x 15 surface, x 150. 46-48, C. macrophylla (Bernardi 9507): 46, aoe side view, x Ps 47, surface, x 50: 48, surface, 00. 1984] DICKISON, CUNONIACEAE 173 PLATE V1 FiGures 49-57. Scanning electron micrographs of seeds of Cunoniaceae. 49, 50, Cunonia schinziana (McKee 23112): 49, side view, x 14; 50, surface, x 40. 51, - pseudoverticillata (McKee 4410), side view, x a 52, 53, Spiraeanthemum katakat (Smith 4371): 52, side view, x 15: 53, surface, x 200. 54, Acsmithia elliptica (Veillon 2844), side view, x 20. 55, 56, A. densiflora Qicsclnan et al. 5222). 55, side view, x 21: 56, surface, x 200. 57, A. reticulata (Galore & Wood, NGF 41005), side view, x 20 PLATE VII FiGures 58-66. Scanning electron oo of seeds of Cunoniaceae, Caldcluvia. 58-60, C. celebica (Havel & Kairo, NGF 84): 58, side view, x 30; 59, surface, x 100; 60, papillae, x 300. 61, C. peeieliee (Hoogland 8538), side view, x 20. 62, 63, C. rosifolia (Walker s.n.): side view, x 50; 63, surface, x 100. 64, C. paniculata (Dombey 713), surface, x 50. 65, C. brassii (Brass 29616), side view, x 23. 66, C. clemensiae (Van Balgooy 969), a x 50. PLATE VIII Ficures 67-75. Scanning electron micrographs of seeds of Cunoniaceae, Caldcluvia. 67, 68, C. paniculosa (Thorne et al. 25938): 67, side view, x 20; 68, surface, x 200. 69, 70, ete (Kalkman 5316): 69, side view, x SO; 70, ee x 500. 71, C. fulva (Van Royen, NGF 18266): 71, oblique side view, x 30; surface, x Eo 73-75, C. nymani (Stevens, NGF 50443): 73, side view, x 15: i surface, x 50; 75, surface, x 500 nN PLATE IX Ficures 76-84. Scanning electron micrographs of seeds of Cunoniaceae. 76-78, Cal- licoma serratifolia (White 10279): 76, oblique side view, x 30; 77, surface, 0; 78, detail of papillae, x 100. 79, 80, Acrophyllum australe (Stauffer et al. 5700): 79, side view, x 50; 80, detail of surface, x 500. 81, 82, Vesselowskya rubifolia (White 11482): 81, side view, x 15; 82, surface, x 150. 83, 84, Weinmannia tannaensis (Ber- nardi 12909): 83, side view, x 30: 84, surface, x 300. PLATE X FIGURES . 93. Scanning electron micrographs of seeds of Cunoniaceae, Weinman- nia. 85, 86, W. racemosa (Philipson et al. 349): 85, oblique side view, x 14; 86, sur- face, x 100. a W. subsessiliflora (Vasquez A130), side view, x 23.88, 89, W. monticola (McKee 23892): 88, side view, x 40; 89, surface, x 400. 90, HW’. pinnata (Sousa & Rico 8118), side view, x 24. 91, W. tinctoria (Bernardi 14516), oblique side view (arrow points to hilum), x 20. 92, 93, W. pullei (Hoogland & Schodde 7242): 92, side view, x 16; 93, surface, x 350. PLATE XI FiGures 94-— ag Scanning electron Sp ae a seeds of Cunoniaceae, Wein- mannia. 94, W. richii (A. C. Smith 6813): , side view, x 40: 95, surface, x 200. 96, ©. alee? (D’Alleizette s. side view, x 20. 97, 98, W. serrata (Ber- nardi 12816): 97, side view, x 40; 98, surface, x 1000. 99, HW” sylvicola (Van Steenis 22314), surface, x 500. 100, 101, W. ae zoniensis (Gacad, FB 27274): 100, side view, 40; 101, surface, x 500. 102, W. glabra (Lansberger s.n.), side view, x 50. 174 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 PLATE XII Ficures 103-11 . Scanning electron micrographs of seeds of Cunoniaceae, Wein- mannia. 103, 104, W. negrosensis (Ramos, Philippine Plants 1287): 103, oblique side iew, x 20; 104, ein x 1000. 105, W. mauritiana (D’Alleizette 2166), side view, x 40. 106, 107, W. descombesiana (Eyma 3590): 106, side view, x 40; 107, surface, x 200. 108, W. trichophora (Pullen 374), side view, x 20. 109-111, W. fraxinea (Fore- man 51): 109, oblique side view, x 20; 110, surface, x 200; 111, surface, x 500. PLATE XIII Figures 112-120. Scanning electron micrographs of seeds of Cunoniaceae, Wein- mannia. 112, 113, W. trichosperma (Zollner 3137): 112, side view, x 40; 113, surface, 0.114, W. vescoi (Van Balgooy 1796), surface, x 200. 115, oa ee . celebica (Van Balgooy 3809): 115, side view, x 40; 116, surface, x 200. 117, W. hutchinsonu (Men- doza, o 18409), surface, x 200. 118, W. purpurea (Kajewski 1738), surface, x 400 119, W. denhamii (Bernardi 13258), surface, x 200. 120, W. decora (Bernardi 11961), nr ae x 300. PLATE XIV Ficures 121-129, sare aeates micrographs of seeds of Cunoniaceae. 121, 12 Codia nitida (Jaffre 2223): 121, side view, x 60; 122, surface, x 200. 123, Pullea ee (Irvine 1253), side view, x 100. 124. P. glabra (Brass 31156), side view, x 100. 125, 126, Pseudoweinmannia lachnocarpa (Clemens 43294): 125, side view (arrows point to arillike outgrowth), x 2; 126, detail of outgrowth, x 75. 127, 128, Platvlophus trifoliatus (Bolus 621): 127, side view, x 50; 128, surface, x 500. 129, Schizomeria whitei(Dickison 205), surface, x 2000 PLATE XV Figures 130-136. Structure of fruits and seeds in Cunoniaceae, transverse sections. 130, 131, Cunonia balansae (Dickison 230): 130, immature fruit; 131, young seed. 132, 133, C. pterophylla (Dickison 238): 132, immature fruit; 133, seed 34. ae robusta (Dickison 248), young seed. 135, Caldcluvia nymanii (Hoogland & Craven 11069), mature seed. 136, Acsmithia densiflora (Dickison 220), seed coat. (C, cotyledon; EN, endosperm: EP, epidermis; IT, inner tegmen: P, papillae: S, ec layer; VB. vascular bundle PLATE XVI Ficures 137-145. Structure of fruits and seeds in Cunoniaceae. 137-140, nape talum dhe bre (Hoogland & Craven 10826): 137, immature fruit, transverse section 38, immature fruit, longitudinal section; 139, hilar end gees seed, transverse section: 140, chalazal end of young seed, transverse section. 141, 142, Schizomeria ovata (Hoog- land 11684), transverse sections: 141, youn g seed; 142, outer portion of seed coat. 143, ieee biglandulosum (Webb 3337), inner region of pericarp, transverse section. 4 Codia nitida (Dickison 221), transverse sections: 144, fruit; 145, pericarp. (CR, rane layer; E, endocarp:; EP, epidermis; IT, inner tegmen; S, sclerenchymatous layer; VB, vascular bundle.) DICKISON, CUNONIACEAE P75 1984] <—f{h AC) PLATE I 176 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 yyy) a / RU ALON OME 4, NG SUSE ANN SII (Vi i Pe. SOY Sp ie it ; OO TEN Aa Pre) i (Oe SS Oy a SRK (sb y vy OCC . 4 { Ki a GUSIe > SS fH wy oe) & -——_1t am PLATE I] DICKISON, CUNONIACEAE 177 1984] CS | | At MS 27 ane Pate II] JOURNAL OF THE ARNOLD ARBORETUM NV I ae \— we ‘ Soma > PLATE IV [VOL. 1984] DICKISON, CUNONIACEAE 179 / OP] PLATE V 180 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 PLATE VI 1984] DICKISON, CUNONIACEAE 181 MY IN./ Ve: fi zea ep = ~ . PLate VII 182 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 ‘ikea PLATE VIII 1984] DICKISON, CUNONIACEAE 183 PLATE IX 184 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 PLATE X 1984] DICKISON, CUNONIACEAE 185 IA a Me UZ 4 f/ a t We | im ME ir GINS ae v7. lq é fs cP] Ae ae 2 | PLATE XI 186 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 PLATE XII DICKISON, CUNONIACEAE 187 1984] PLATE XIII JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 188 LATE XIV P DICKISON, CUNONIACEAE 189 1984] PLATE XV 190 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 65 St ae zi; ¥ wy Pitre, Aes at, ae oie Eta vn seer ys wa PLATE XVI 1984] TOMLINSON & VINCENT, RHAPIS EXCELSA, X Lee ANATOMY OF THE PALM RHAPIS EXCELSA, X. DIFFERENTIATION OF STEM CONDUCTING TISSUE P. B. TOMLINSON AND J. R. VINCENT DESPITE THE APPARENT VASCULAR COMPLEXITY of the palm, its vascular de- velopment, structure, and function can be perceived quite readily. Compared with branched trees, it is architecturally simple (Hallé ef a/., 1978). Metaxylem and protoxylem are structurally and topographically very distinct. Vascular tissues are radially as well as tangentially separated (unlike those of dicotyle- dons), and due to the relatively massive meristematic region and the long time course of differentiation, successive events are widely separated in time as well as in space. Finally, primary structures are not obscured by later development of secondary vascular tissues. In dicotyledons recent advances in our under- standing of vascular developmental processes have required very precise anal- yses of serial thin sections, as in the work of Larson (1982; and earlier papers cited therein) on Populus. In this paper we consider the sequence of initiation of xylem and phloem within the procambial template and the changes within these tissues as both radial and axial extension take place. Special attention is given to xylem dif- ferentiation in traces to a given leaf at various stages of its development, a topic being investigated physiologically by John Sperry at Harvard Forest. In the first paper in this series (Zimmermann & Tomlinson, 1965), a cine- matographic method of analysis was used to describe the course of vascular bundles in the mature aerial stem of the small palm RAapis excelsa (Thunb.) Henry. In a topographic sense, traces serving a given leaf (leaf traces) usually branch to give an axial bundle that becomes a leaf trace at a higher level. Axial bundles in the region of leaf-trace departure are connected by short bridge bundles. In addition to establishing the principle of vascular continuity in the stem, that paper demonstrated the changes that take place in individual mature bundles throughout their length. It provided a basic, general framework (since termed the “Rhapis-principle’’—see Zimmermann & Tomlinson, 1972; Tom- linson, 1983) for understanding the vasculature of monocotyledonous stems, which may be described as a regular pattern of outgoing leaf traces branching to generate topographically axial bundles and other derivatives. To add to the topographic analysis, in the fourth paper of the series (Zimmermann & Tom- linson, 1967), the sequence of initiation of strands within the developing crown was analyzed in terms of their inception as procambial strands. The principle of vascular development that has been shown to be generally applicable to monocotyledons was demonstrated. Procambial strands that connect to young leaf primordia are initiated within a cap of meristematic tissue. They are © President and Fellows of Harvard College, 1984. Journal of the Arnold Arboretum 65: 191-214. April, 1984. 192 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 continuous with an existing leaf trace but are at first uncommitted in a distal direction to a specific future leaf primordium. These processes are difficult to visualize in static diagrams but are shown clearly in recently produced dem- onstration films (Zimmermann & Mattmilller, 1982). We believe that the basic nature of the Rhapis type of construction has wide relevance to an understanding of monocotyledonous vascular development (Tomlinson, 1983). Our overall objective is to present an understanding of the palm crown in terms of anatomy, development, and physiology. METHODS The analysis involved two successive steps (see FIGURE |) because the de- veloping region of the shoot (the crown) is an extended structure. First, the sequence of development of vascular tissue within individual bundles was studied at early stages of development (see FiGure 3). In this phase radial expansion predominates. Second, a less-detailed analysis was carried out of late stages of development; dye-infusion techniques were used and unembedded material of long apical portions was sectioned (see FiGure 4). In this phase longitudinal expansion predominates. Although technically necessary, such separation into phases 1s artificial since growth 1s a continuous process. The first method provides information about the development of the vascular connection, via protoxylem, between stems and young leaves until initiation of predominant axial extension of the internode below a leaf with an expanded blade. This initial analysis can be done with stem pieces (including the shoot apex) up to a length of about 1.5 cm (FiGure |, A). The second method gives information about the establishment of connection between the leaf and the metaxylem of the stem during the period when the leaf blade is fully expanded but the intercalary extension of the leaf sheath is still incomplete (FIGURE |, B). Final maturation of metaxylem occurs as soon as internodal extension is complete. Again, separation into phases of development is artificial because growth is continuous, but it is important to appreciate that blade expansion precedes maturation of the leaf sheath and its associated internode. The “‘first fully mature leaf’ is therefore the first leaf in which extension of all these regions is complete. The stage at which a given developmental event occurs varies by as much as three plastochrones, depending upon the size of the crown. A leaf usually first becomes visible externally somewhere between plastochrones P8 and P11 (P| is the first microscopically visible leaf primordium). Internodal extension is generally complete somewhere between P14 and P17. For purposes of de- scribing the position of leaves that are only visible externally, the youngest visible leaf may be referred to as leaf 1, the next youngest as leaf 2, etc. Without dissection the numbering of such leaves by the plastochrone (P) system cannot be done. In the crown analyzed we recognized about seven leaf primordia enclosed within the apical bud and not visible externally (as in FIGURE 5, where the position of P8 corresponds to the “spear leaf’’—1.e., the first externally visible leaf in the crown, its blade still unexpanded). 1984] TOMLINSON & VINCENT, RHAPIS EXCELSA, X 193 crown a I, first __4° expanded (ho leaf blade Vv) © O }: 3 Cc f © E oO £ ne) TZ x be LA. as vO first o extended = internode A g £ 6 TO o ue) Cc he ® 7 6) 0.5cm Figure |. Rhapis excelsa, diagrams showing size of specimens used to analyze vas- cular development in shoot (2 methods): A, camera lucida drawing of median longitudinal section of apex about same size as that used in analytical plots (see FiGure 5); B, diagram of axis used in experimental dye injection of single leaf, foreshortened 50%. Numbers refer approximately to corresponding expanded leaves in visible crown. 194 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 PLOTTING OF EARLY STAGES To plot the early stages of vascular bundles, we examined three section series, including one described in a previous paper (Zimmermann & Tomlinson, 1967). These series were derived from apices from which all but the youngest enclosed leaves were trimmed. They were fixed, embedded, sectioned at 8-15 um, stained in safranin and Delafield’s haematoxylin, and mounted in the usual way, but with only a single section on each slide. Plotting was carried out by the drawing and shuttle methods of Zimmermann and Tomlinson (1966)— i.e., either with a drawing attachment to a Wild M-20 microscope or with two separate microscopes connected by an optical bridge. These methods provided precise alignment for successive sections. Single major leaf traces belonging to successively older leaves were followed, starting with Pl (the youngest leaf primordium), in a basipetal direction from the level of insertion of the leaf. In Rhapis the crown is a shallow cone (Figure |, A; see also Zimmermann & Tomlinson, 1967, fig. 4), so it is possible to follow traces continuously in one direction. Positions of each trace measured at regular intervals were plotted on graph paper, with only the radial coordinate considered, the helical or tangential displacement being _ for diagrammatic purposes (see Zim- mermann & Tomlinson, 1967, fig. At each plotted level the number See xylem elements was count- ed. To facilitate recognition of differentiated protoxylem elements, transverse sections were viewed through partly crossed polarizing filters; the birefringence of cell walls of fully mature elements rendered them conspicuous. It was con- siderably more difficult to recognize protophloem elements at very early stages of vascular differentiation in bundles that were cut somewhat obliquely. The criteria that we used were presence of cell walls densely stained with haema- toxylin, and cell contents relatively unstained or even clear. Sometimes simple sieve plates could be seen in face view, making the identification of sieve elements unequivocal. The principal objective was to establish whether or not phloem was continuously acropetal in its differentiation. GENERAL ANALYSIS OF LATE STAGES To show late stages of xylem differentiation and particularly to identify the vascular contribution of a given leaf in its final stages of development, dye ascent and descent experiments were conducted using Schiff’s reagent (reduced- acid fuchsin) after a preliminary perfusion with 0.5 percent periodic acid as an aldehydic mordant. This was similar to the method used by Priestley and colleagues (1935) to trace bundles from specific leaves in Alstroemeria with Magdala red as a marker stain but without any mordant. In the ascent exper- iment the stem was cut about 20 cm below the shoot apex. The cut end was immersed in dye, and suction was applied to a cut leaf. Distally, dye moved preferentially into the traces of the leaf to which suction was applied. In the more-informative descent experiments, a flask containing the dye was attached via an air-tight seal to the cut petiolar stump of a given expanded leaf of a detached shoot. Dye moved only into the traces of the injected leaf. On different shoots successive leaves from the youngest to the fifth-youngest expanded 1984] TOMLINSON & VINCENT, RHAPIS EXCELSA, X 195 leaves were infiltrated. The dye penetrated the shoot over a period of about 24 hours: it could then be recognized in its colored, oxidized form in the cell walls of the leaf-trace xylem continuous from the cut leaf, as seen in sections cut at successively lower levels (e.g., FiGuRE 4). In this way the trace system from a given leaf could be recognized in freehand sections of the shoot at successively lower levels. Sections were dehydrated and mounted in Permount without further staining. Traces were followed to a maximum distance of 36 cm—i.e., up to about 15 internodes below the morphological level of insertion of the injected leaf. Additional sections cut at intervals from other shoots, stained in phloroglucinol and concentrated HCl, showed the general progression of late stages of maturation of lignified tracheary elements, but without reference to a particular leaf. These sections were useful for recognizing the highest level of maturation of metaxylem. RESULTS GENERAL FEATURES OF VASCULAR DIFFERENTIATION EARLY STAGES (RADIAL EXPANSION PREDOMINANT). Observation of single sec- tions gives general information about the progress of vascular differentiation at a single level. Ficure 2 represents the appearance of a vascular bundle at an approximately comparable level in successive plastochrones, so that on- togenetic events in a single bundle are simulated. The illustrations were pro- duced by selecting an arbitrary standard level (a major leaf trace at its most central location) and, with the aid of the drawing apparatus, making a drawing or a comparable bundle serving a leaf at successive plastochrones. (“Standard level” refers to an equivalent position below successive older leaves.) The drawings are thus illustrative of the general process of vascular differentiation at that particular level. Other sequences at other levels are shown photograph- ically in FiGure 3. Precise information about developmental events in different parts of representative individual bundles is included in the plots (FIGURES 5, 6). The procambial strand is visible in transverse section as a group of narrow elements that retain a meristematic appearance in contrast to the surrounding vacuolated cells (FiGurEs 2, A; 3, A). We have not been concerned in this study with the details of the appearance of procambial cells or with the method by which the diameter of the vascular strand is increased, although this is an important early process in the development of the vascular bundle (e.g., the changes within Ficures 2, A-E, and 3, A, B). Cell divisions in later stages of procambial development are mainly longitudinal. In longitudinal section the strand comes to have a characteristic “tiger-tail’” appearance because nuclei of adjacent, recently divided procambial cells lie at the same level. Cells in the center of the bundle vacuolate early, in contrast to those in the inner (xylic) and outer (phloic) regions, but cell division still continues in them and produces a temporary ‘“‘cambiumlike”’ condition seen in transverse section (FIGURE At the standard level illustrated (FiGuRE 2), the first appearance of vascular JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 196 aes Cie=28C5O OC (a Kies laste] TRS ae 19480 eae, x \ i rh 280 ee ecenste- SS sasese ss, tee or eater , = Oe Bt REO TOV A va BIC laa SES las O as 20Sse3. Ty LE EE EF OY SS 2s tS A ) E he ® at Ss t TAL EA) EA ase sreceuNen tars cM AOS e X) ORLOV STA AR, OF aes é Sap ae: ae & vedas JESSY eecee, ne @ es Se \Y MOO¢d Rhapis excelsa, transverse sections of developing leaf traces to progres- FIGURE 2. 2.8mm (P11), differentiating metaxylem vessels G, | differentiation of metaxylem (mxy.); 1984] TOMLINSON & VINCENT, RHAPIS EXCELSA, X Log tissue is the development of protophloem toward the outside (peripherally oriented) part of the procambial strand (Ficure 2, B). In the permanent prep- arations used in this analysis, differentiated protophloem sieve elements have somewhat thickened angular walls. These walls stain densely with haematoxy- lin, and either the protoplast stains lightly or the cell lumen appears completely empty. Simple transverse sieve plates are often evident. Protophloem elements of this kind have a diameter of about 8-10 um. At the given level sieve-element (phloem) differentiation scarcely precedes protoxylem differentiation and in some strands even follows it (FiGURE 2, C). We found no precise evidence for discontinuous sieve-element differentiation, although in a few bundles uncer- tainty in recognizing phloem elements sometimes gave this impression within a few adjacent sections. Later differentiation of sieve elements at any one level was always in a centripetal direction and involved elements somewhat wider (10-15 wm) than those first detected (FiGuRE 3, B). In association with these changes, there was evidence of disruption and collapse of the youngest protophloem elements. In transverse appearance the lumen of the first-formed sieve elements becomes irregular and is finally occluded by expansion of surrounding cells. In still later stages the position of the protophloem is marked by a densely stained region of cell-wall material. In the axial portion of a mature bundle, the protophloem region is obscure and included within the innermost bundle-sheath fibers. It proved difficult in the kind of sections used to quantify (in terms of numbers of functional sieve tubes) the sequence of events involved in early phloem differentiation because of the problems of recognizing both early stages of sieve- element differentiation and their later collapse. In transverse sections of young leaf traces, protoxylem can first be recognized unequivocally as enlarged cells (ca. 10-15 wm diameter) with thick, lignified walls that stain with safranin (FicurE 2, C, D). Polarizing optics confirmed the existence of a thickened birefringent cell wall. In addition the cell lumen always appears empty. The first protoxylem element occurs on the inner side of the procambial strand, remote from the protophloem, and is followed by a succes- sion of elements that appear in centrifugal order. Later-formed elements are somewhat wider and more conspicuous; they form a circular or wedge-shaped group of cells in transverse section (FIGURE 2, G). The intervening area between protoxylem and protophloem is occupied by procambial cells, and differentia- tion within this region varies depending on the level at which one examines a given strand, as is indicated later (FicureE 3, C, D). The distance between first- formed xylem and phloem elements actually increases as the bundle matures, due to the increase in number of procambial cells by the continued longitudinal divisions already mentioned. We have not followed developmental changes in detail in longitudinal sec- near maximum diameter. (Black circles = nuclei, stippled areas = sieve tubes of pro- tophloem, solid black walls = protoxylem, lumen outline dotted = differentiating pro- toxylem and metaxylem elements.) 198 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Y" MT eT o ere aA al Sat a ie mi <) AL Ve eS: “AS mis A ark : cite DERN) St eRe Mipey es. Bases ia FP qe Ce Nea tt ry < 4 ; Cit oP Deeg eRe: - Ps Mower oa. oe WHATS MK et eaane ot Na ers > ; frat es Cure ea er ee A 425 eos Sah a A c Raat . isa ‘ . a att o “ae Basan i's RNS ata’ he, 1.00 . x ‘ «®, Be \) wy ee SS reonte: yes! ‘ ee < * EL yee ar SS, en Be 4x) Tt Neues: Dre Ts peg? FiGurE3. Rhapis excelsa, photomicrographs of 15-um-thick transverse sections from paraffin-embedded material. A, 0.3 mm below shoot apex: leaf-trace system to Pl, P2, and P3 still discretely recognizable within stem, trace system to P4 within leaf base. B, 1.8 mm below shoot apex: major leaf trace to P8; protoxylem and protophloem well developed, early metaxylem differentiation evident. C, 5.7 mm below shoot apex: major leaf trace to P10 close to periphery of central cylinder; protoxylem and protophloem still incompletely differentiated (no metaxylem in this part of leaf trace). D, 15 mm below shoot apex: major leaf trace; protoxylem almost completely differentiated, metaxylem elements not in contact with protoxy] l t pletely expanded but still thin walled. E, 13.8 mm below shoot apex: axial bundle; single incompletely differentiated metaxylem 1984] TOMLINSON & VINCENT, RHAPIS EXCELSA, X 199 tions. Serial sections show that the protoxylem elements are short, but with annular or helical wall thickenings, and are thus presumably extensible. Pro- tophloem elements are also short. At the standard level illustrated in Figure 2, metaxylem differentiation begins at a very early stage when there is little mature protoxylem (see FIGURE 3, B). In transverse view the first evidence for metaxylem differentiation is the enlargement and vacuolation of a pair of cells toward the middle of the bundle but laterally widely separated from each other (Ficure 2, F). These initials subsequently widen considerably and block out the characteristic metaxylem pattern. The walls of the enlarging vacuolating cells remain unthickened and contrast with the lignified, thick, birefringent walls of the protoxylem (FIGURE 2, G). Maturation of metaxylem occurs very late, well below the crown, and only in bundles that are directly in association with older expanded leaves and that run through fully extended internodes (i.e., in the unshaded part of FIGURE 1, B), as is described later. Depending on the level examined, protoxylem may or may not be in direct lateral contact with the metaxylem (cf. Ficure 3, D and E); in the distal part of any axial bundle, at the level of its divergence toward a leaf, protoxylem is separated from metaxylem by a broad region of procambial cells that mostly mature as parenchyma cells. In late stages of differentiation, the cells of this conjunctive parenchyma become somewhat radially seriated (FIGURE 2D). indicative of further late, regular tangential division, which is completed before metaxylem differentiation begins. Evidence for the time and regularity of di- vision is provided by the identical length and the coincident end walls, at any one level, of the wide vessel elements and the metaphloem sieve tubes (Par- thasarathy & Tomlinson, 1967). However, the situation is complicated by the late differentiation of tracheary elements on the outer face of the protoxylem, as is described later. Fiber differentiation outside the protophloem is initiated early but is long continued. Maturation of fibers begins in the region of the protophloem, but the pattern within the future fibrous sheath of the vascular bundle becomes complex. The extent of apical intrusive growth of differentiating fibers is not known. However, in larger palms there are regular trends of change of fiber length throughout single stems, indicating a high degree of endogenous control of fiber length but some correlation with internode length (Tomlinson & Zim- mermann, 1967). LATE CHANGES (INTERNODAL EXTENSION PREDOMINANT). In any one leaf trace followed basipetally, the gap between protoxylem and metaxylem decreases, and at some level incompletely differentiated protoxylem elements are seen to be contiguous with incompletely differentiated metaxylem elements (FIGURE 3, E). Fully differentiated contiguous protoxylem and metaxylem elements vessel contiguous with incompletely differentiated (presumed) protoxylem elements. (Empty squares = differentiating protoxylem, solid squares = mature ee solid stars = differentiating metaxylem. Scale for A = 50 um, for B-E = 10 200 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Figure 4. Rhapis excelsa, transverse sections about 50 um thick from fresh, unembedded material after dye-descent injection experiments. Dark staining indicates presence of Schiff’s reagent in xylem. A, major leaf trace to first fully expanded leaf 3.9 cm below its morphological insertion: protoxylem and ) t and closely contiguous so dye moves from former to latter (downward in this experiment B, major leaf trace to second fully expanded leaf 3 cm below its morphological insertion: protoxylem fully differentiated and conducts dye readily downward; metaxylem incom- pletely differentiated and not contiguous with protoxylem so dye not conducted. C, major 1984] TOMLINSON & VINCENT, RHAPIS EXCELSA, X 201 occur only in internodes in which extension is complete (FiGure 4, A). Trans- port of water is then possible between the two types of element, as the dye descent and ascent experiments show. In axial bundles a single metaxylem element is differentiated in the absence of any protoxylem. This construction also applies to bridge bundles (and presumably to branch traces, although we have not examined their development in detail). The peripheral part of each bundle differentiates as the fibrous cap, the extent of which is directly related to the proximity of the bundle to the stem center: peripheral bundles develop a wide fibrous sheath, central bundles a narrow one. Stegmata (silica cells) are conspicuous in early stages of maturation of the fibers (e.g., S in FiGureE 4, B). Although we have not followed individual bundles directly over long dis- tances. the overall maturation of conducting elements can be seen in sections at progressively lower levels. In developmental terms there is thus an advancing “front” of maturation so that overall vascular differentiation is at about the same level of advancement in all bundles. VASCULAR DIFFERENTIATION IN THREE DIMENSIONS Changes in vascular pattern with bundle position and age may first be ex- amined from plots of major bundles to leaves at successive plastochrone in- tervals. These positions are plotted collectively in FiGure 5, with details of the apical region enlarged in Figure 6. We should emphasize that quantitative information relates to the one crown from which the plot was made. Different crowns give different absolute values, but the relative values are much the same. The differences (largely quantitative) between intermediate and minor bundles are explained later. This reductionist approach demonstrates the basic pattern for all bundles, but is free of the topographic complexity occurring in the crown due to the large number of bundles involved. It should be reemphasized that, according to the earlier study of Zimmermann and Tomlinson (1967), differentiation of the axial bundle is continuously acropetal in relation to the meristematic cap just below the shoot apex. Evidence that some vascular tissue develops in a continuous acropetal direction comes from the observation that undifferen- tiated vascular tissue is continuous below with differentiating and ultimately leaf trace to fifth fully expanded leaf 2 cm below its morphological insertion: protoxylem and metaxylem both fully mature but only protoxylem conducts dye downward, meta- lower level; hydraulic constriction apparent). (Open star = mature metaxylem vessel contiguous with protoxylem; solid triangles = protoxylem with densely stained walls; solid circles = metaxylem not in contact with protoxylem, unstained in this experiment; solid stars = differentiating and nonconducting metaxylem; S = stegmata. Scale bar = m.) X3adV MO1S8 WW JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 MM FROM STEM CENTER 2 3 dfdmxy — Rhapis excelsa: schematic longitudinal view of distribution of differen- Pe oe tissues within major bundles to several successive leaves in developing crow me series used to produce FiGure 2 (see FiGure 6 for apical details). Axial dimension foreshortened about 50%. All leaves shown as if occupying single or- thostichy instead of actual 2/5 spiral phyllotaxy; internal helical course of vascular bundles also ignored. Irregular width of leaf bases an artifact of plotting and display. 1984] TOMLINSON & VINCENT, RHAPIS EXCELSA, X 203 mature vascular tissue. These statements apply to protophloem and metaxylem, which are continuous acropetally. However, protoxylem is discontinuous and differentiates bidirectionally. PROTOPHLOEM DIFFERENTIATION. Within the median leaf trace to Pl, proto- phloem is not recognizable until nearly 1 mm below the shoot apex has been traversed (FiGureE 6). This is true of median major traces to the next four older leaf primordia (P2—P5). The precise level is somewhat uncertain because It is not easy to detect the first-formed protophloem element. Once located, how- ever, protophloem can be traced continuously in a basipetal direction in the axial bundle. From sections of older vascular bundles, it seems that the dif- ferentiation of phloem, once initiated, is continuous at any one level, with new elements added in a centripetal direction (FicuRE 2, D-G). The distinction between elements differentiated during (protophloem) and after (metaphloem) elongation of the vascular procambium is somewhat arbitrary, but can be inferred indirectly by considering the level at which metaxylem first matures (which indicates cessation of organ extension). We have not investigated struc- tural differences between protophloem and metaphloem established by this indirect method. rotophloem continuity into a leaf is first evident in a trace to P6 and is then found in major traces to all older leaves (FiGURE 5). PROTOXYLEM DIFFERENTIATION. Protoxylem was not detected at any level in a major trace to the youngest leaf primordium (P1) within the sectioned series. It can be seen in a trace to P2, but only at a considerable distance below the shoot apex—about 4 mm in an axial direction (FIGURE 5, upper px). Traced further in a basipetal direction, this protoxylem disappears at a level about 16 mm below the shoot apex (FiGuRE 5, lower px). This is the developmental origin for the basipetal discontinuity of the protoxylem, evident in the mature stem. Since within a single plastochrone (P1 to P2) protoxylem appears and becomes elaborated over a distance of 12 mm, its differentiation is obviously very rapid. Further extension of this distance until protoxylem is continuous into the leaf base does not occur until the leaf is in position P6. In this leaf there is initially a short discontinuity within the leaf base (between px and px in Ficure 6) representing the level of the intercalary leaf meristem; protoxylem can be seen in the leaf base above this level of discontinuity in the bundle investigated. Protoxylem continuity between leaf and stem is established in P7 wa Single major trace plotted to P1, 2, 4, 6, 8, 10, 12, and 13. In this crown P8 is youngest externally visible leaf (“‘spear leaf’); P9, first fully expanded leaf; P14 or P15 (outside this view), youngest leaf inserted above fully extended (mature) internode (see FIGURE 1). (Dashed lines = procambium (proc.), with or without differentiated vascular tissues; hatched lines = mature protophloem (prtphl.); dotted lines = mature protoxylem (prtxy.); solid lines = vascular bundle with differentiated (but still immature) metaxylem (dfdmxy.); AB = level of departure of axial bundle from leaf trace; px (above and below) = upper and lower limits of differentiated protoxylem in axial portion of major leaf trace to P2; arrows = bundles diverging from outgoing leaf trace with further extremity not plotted.) 204 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 MM FROM STEM CENTER Tey YLT wey thts Sa Me XddV MO1S¢E WW MO l Ficure 6. Rhapis excelsa, enlargement of crown region of FiGure 5 to show early stages of vascular differentiation in major leaf trace to P1, 2, 4, 6, and 8. (Arrowheads = most distal level at which metaxylem differentiation first detected: discontinuity px to px in trace to P6 = discontinuity of mature protoxylem in leaf base. Shading conventions as for FiGure 5.) (not plotted in Ficures 5 and 6) but is represented within the vascular bundle only by a single file of protoxylem elements at the leaf base. All older leaves (e.g., P8 in FiGure 6) include protoxylem continuous from stem to leaf and in several overlapping files. This information demonstrates the intrinsic discon- tinuity of protoxylem. From the change in topography of a given bundle as a result of combined radial and axial displacement, it is reasonable to describe the further overall course of protoxylem differentiation as “bidirectional.” At any one level the amount of protoxylem progressively increases because differentiation of new elements proceeds faster than old elements are obliterated (cf. Figures 2, A and B; 3, B and C). Because of the protoxylem discontinuity, there is a basal decline in number of protoxylem elements that can be inter- preted as evidence of the basipetal component of bidirectional differentiation. Ficure 7, a plot of the total number of elements in all vascular bundles in the 1984] | TOMLINSON & VINCENT, RHAPIS EXCELSA, X 205 1000 5 “spear ° | leaf” 100 - 14 8 8 ° Total mature pxy elements Vbs with mature pxy ae f 2 6 8 10 12. eine NO. Figure 7. Rhapis excelsa, proton yicm development at leaf insertion of successively primordia (P1 is youngest microscopically visible primordium); open circles = total ma- ture protoxylem elements at same level.) P9 is first leaf with fully expanded blade; protoxylem maturation continues in leaf base for at least 4 plastochrones after leaf blade has expande base of each leaf at successive plastochrones, indicates the extent of protoxylem connection of successive leaves but underestimates (because of protoxylem obliteration) total protoxylem produced. The information is sufficient to show that P8, the “spear leaf,” is the first leaf with appreciable xylem contact with the stem, but that later leaves add considerably to their xylem transport capacity without adding new vascular bundles. This diagram can be compared with fig. 4 in Zimmermann and Tomlinson (1967), which shows the total of all vascular bundles in successively older leaves. METAXYLEM DIFFERENTIATION. Two stages of metaxylem differentiation can be recognized: early, when the conspicuous metaxylem vessels are first initiated (e.g., FiGuRES 2, 3, 6); and late, when the cell contents are lost and wall thick- ening is completed (unshaded portion of FiGure 1). In the mature vascular bundle at distal levels close to the level of branching as a leaf trace (e.g., lower 206 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 AB in Ficure 5), the conspicuous metaxylem is represented by either a single vessel or a widely separated pair of vessels. Overlapping ends of vessels are commonly seen, especially in the region of attachment of bridges and branch traces (e.g., left-hand group of vessels in FiGurE 2, F). At any level where metaxylem is present, the elements are first visible as cells that vacuolate and enlarge conspicuously, in contrast to adjacent cells, most of which mature as narrow conjunctive parenchyma cells. This contrast occurs, for example, in a major trace to P| at an axial distance of about 1.5 mm below the shoot apex (Ficure 5). Significantly, this is well before protoxylem is differentiated in the same strand at this level. Protoxylem in the leaf trace does not occur much in advance of the differentiating metaxylem ‘until about the time when the as- sociated leaf is in position PS. After this, protoxylem is always continuous into the leaf base, as we have established. Metaxylem does not differentiate into the outgoing leaf trace but is continuous axially via bridges to adjacent axial bundles. These axial bundles themselves become recognizable as procambial strands in the crown just below the meristematic cap (Zimmermann & Tom- linson, 1966). However, bridges with early stages of metaxylem differentiation do not become recognizable until about the level of insertion of P12—1.e., almost 5 mm below the shoot apex proper in an axial direction. Such a leaf ts the fourth-youngest fully expanded leaf in the crown illustrated in Ficure 5. The relatively rapid advance of metaxylem differentiation within an axial bun- dle can be seen by comparing the procambial strand diverging from the leaf trace to P10 at the upper AB with that diverging to P12 at the middle AB in FiGureE 5. Late stages of metaxylem maturation involve development of secondary wall layers and their lignification, together with loss of cell contents. Examination of free-hand sections stained with phloroglucinol and concentrated HCI shows lignified walls in wide metaxylem vessels at linear distances of about 3-4 cm below the shoot apex. This is about the level of insertion of the fifth leaf below the first fully expanded leaf and is always at the level where internode elongation has just ceased (FiGureE |, B, unhatched portion). Maturation of metaxylem appears to be continuously acropetal. In addition to the wide metaxylem, there are late-differentiated tracheary elements on the outer surface of the protoxylem strand that may not be totally mature until the internode (and therefore the vascular bundle) is completely extended (Figure 3, D, E, open squares). This becomes obvious when late stages of vascular differentiation are examined, but it was not considered when the mature structure was described. Consequently, 1t was assumed that all narrow elements on the inner face of the leaf trace were protoxylem tracheids (Zimmermann & Tomlinson, 1965). The late-maturing elements in this po- sition, however, must be metaxylem, and they possibly correspond to some of the narrow vessel elements observed by Zimmermann and Sperry (1983) in macerated material from stems. They occupy only a limited part of the leaf trace since they occur just below the level where the protoxylem first diverges from the metaxylem within a given bundle. However, they are functionally important because this is the region in which interchange of water between metaxylem and protoxylem occurs (FIGURE 4, A). 1984] TOMLINSON & VINCENT, RHAPIS EXCELSA, X 207 ESTABLISHMENT OF THE VASCULAR CONNECTION The level at which the “descending” protoxylem of the leaf trace becomes contiguous with the ‘‘ascending” mature metaxylem can be determined from dye descent experiments, since the downward-moving dye initially descends in the protoxylem and can only enter the metaxylem of the axial bundle when a) this is mature, and b) the two tissues are in direct contact (FIGURE 4). Because the dye is initially restricted to the traces connecting the injected leaf to the stem, the vascular supply to that leaf can be identified in the stem. The Jux- taposition of the two types of xylem can also be readily recognized in sections. Movement of dye from protoxylem to axial metaxylem occurs (see FIGURE 4, A) in the trace system of leaf P9 and can be observed in the internode below P13 or P14—i.e., the internode in which extension has just been completed. (This association is, of course, rational and a simple consequence of the defi- nition of metaxylem, which cannot mature completely within an extending organ.) The area of overlap (vascular insertion) is also mature at this level. At a higher level in the same bundle, only protoxylem 1s fully mature and conducts the dye; metaxylem is still undifferentiated (see FiGure 4, B) and cannot trans- mit the dye even if it is contiguous with the mature protoxylem. The same considerations apply when protoxylem and metaxylem are not contiguous, regardless of whether metaxylem is mature (see FiGure 4, C) or not (see FIGURE D ’ MATURATION OF INTERMEDIATE AND MINOR BUNDLES Each leaf is supplied with about 100 vascular leaf traces that connect with the central cylinder of the stem. These have been termed major, intermediate, and minor bundles according to their topographic arrangement and time of appearance in the leaf base (Zimmermann & Tomlinson, 1965). Major bundles appear earliest and diverge from the stem center; intermediate ones appear later and diverge from a position nearer the stem periphery; and minor ones appear last and are restricted entirely to the periphery of the central cylinder. These designations are arbitrary since there is a developmental continuum (Zimmermann & Tomlinson, 1967). There are corresponding differences in the leaf-contact distance, with major bundles many internodes long, and minor bundles only a few. In addition to the 100 vascular leaf traces, there are about 1000 cortical bundles. These appear last and, by definition, diverge only from the stem cortex. They are discontinuous basipetally or anastomose among themselves. The “rules” that govern vascular development in major bundles also apply to later-formed bundles, except that maturation of cell types occurs later. Dif- ferentiation of vascular tissues in progressively more peripheral (minor) bundles takes place during shorter periods of extension; consequently, minor leaf traces have very little protoxylem. Metaxylem matures somewhat later than in major bundles but according to the same principles. Although it would be possible to quantify these statements with the techniques available, no fundamentally novel information would be obtained. 208 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 DISCUSSION The information presented above, when considered together with our pre- vious studies on the course and structure of vascular bundles in mature stems and the process of initiation of vascular bundles themselves, allows us to present a highly integrated view of vascular development in the palm crown. Vascular bundles establish continuity between stem and leaf as procambial strands and increase in number at the level of leaf insertion. When a bundle makes contact with a leaf in this way, protophloem and metaxylem elements are already differentiated in the lower part of its course; the differentiation and maturation of these tissues is continuous acropetally, but metaxylem matures in such a way that it diverges into bridges or branch traces and does not continue along the leaf trace proper into the leaf base. Protoxylem first appears in the distal part of the trace when the associated leaf is in position P2. It is discontinuous both distally into the leaf and proxi- mally into the axial portion of the leaf trace. Protoxylem continuous from stem to leaf first occurs when the leaf is at about position P7. Metaxylem does not begin to mature until the subtending internode of a given leaf has ceased to elongate. However, younger leaves have an indirect vascular continuity with the axial metaxylem because mature protoxylem and metaxylem are contiguous in completely mature internodes, with the mature protoxylem distally continuous through actively extending tissues and into the leaf itself (FiGuReE 8). Xylem contact of about four or five leaves with expanded blades is made through tissue extension in this way. This means that the xylem- to-xylem contact between stem and leaf is still developing after the leaf blade has expanded. Sperry (pers. comm.) has preliminary information showing that this change affects the transpirational capacity of the leaf in its early life as an expanded organ. HYDRAULIC CONSIDERATIONS Transport capacity in the xylem to the developing leaf is indicated approx- imately by Ficure 7 as the total number of protoxylem elements in the leaf base at successive plastochrones, which is at first dependent on—and then subsequently independent of—the total number of vascular bundles that dif- ferentiate in the leaf base. The measure is incomplete and would be better indicated by the sum of the fourth power of the diameter of all elements Zimmermann & Sperry, 1983). The basipetal connection of this protoxylem is dependent on the overlap between protoxylem and metaxylem at lower levels. The vertical distance over which metaxylem and protoxylem are directly contiguous within any one vascular bundle is limited. It may be termed the vascular insertion of the leaf trace onto the axial system (Sperry, pers. comm.), in contrast to the morphological insertion of the leaf base at the node itself. Only in the region of overlap can water move directly from metaxylem into the protoxylem of leaf traces; this, therefore, is the significant level of leaf insertion as far as water transport is concerned. However, vascular continuity between axial and leaf-trace xylem is established very late in crown develop- ment, in internodes in which extension has just been completed. It is dependent 1984] TOMLINSON & VINCENT, RHAPIS EXCELSA, X 209 ~™N l we 1 Pa Ficure 8. Rhapis excelsa, fundamental aspects of xylem development and topog- raphy. Left: develop tal relationships between } mbium (dotted lines), protoxylem (hollow circles), and differentiating metaxylem (solid line) in crown. (Numbers = leaf positions, topographic relationships schematic—see Ficures 5, 6.) Right: relationship etween protoxylem (hollow circles) and metaxylem (solid line) in mature stem; 2 com- plete leaf contacts of major stem vascular bundle. (A = “foliar’’ component of xylem m,; continuous into leaf; C = region of overlap between A and B where mature protoxylem and metaxylem in direct contact (region of “‘vascular” insertion within stem, in contrast to “morphological” insertion of leaf at node).) on elements differentiating late as protoxylem, next to metaxylem elements in which maturation has just been completed. Distally, water moves into leaves only through protoxylem, as in the traces to P8, 10, 12, and 13 in Ficure 5. Major bundles that have extensive protoxylem connection with the leaf base continue to develop new tracheary elements between first-formed protoxylem and still-immature metaxylem within extending regions. Over the region of 210 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 65 vascular insertion (i.e., where protoxylem and metaxylem are laterally contig- uous), differentiation of protoxylem continues from initials adjacent to im- mature metaxylem elements. The water-transport pathway is only completed when metaxylem finally matures. Dissection of a number of crowns showed that there are four or five incom- pletely mature leaves (in which the leaf blade is expanded but the base is still immature) older than the spear leaf. The internodes below the node of insertion of each of these leaves is also still extending. Consequently, the first fully mature leaf associated with the first fully mature internode is about the fifth below the spear leaf and about P13 in this crown as a whole. All internodes below this level are completely extended and support fully mature leaves. The total num- ber of leaves in the crown varies according to the position of the crown: fully exposed crowns include fewer visible and enclosed leaves (about nine of each), while those in the shade support more visible and enclosed leaves (about twelve ofeach). This range of quantitative variation has to be considered when absolute statements are made about the level of vascular differentiation below the shoot apex. Time of maturation of tissues may vary by as much as three plastochrones in different crowns. Nevertheless, similar conditions determine the develop- ment of the hydraulic connection established in each leaf as it develops. COMPARISON WITH OTHER MONOCOTYLEDONS The most directly comparable study is that by Esau (1943) of the ontogeny of the vascular bundle in Zea mays. Although there are some common features, there are also appreciable divergences in the process of vascular bundle de- velopment between this grass and the palm studied here because of differences between them in size and general organization. A particular difference is in the axial meristem: it is interrupted in the stem of corn, and uninterrupted in the palm (see Fisher & French, 1976). The corn stem thus has a nodal plexus that is absent from the palm. Consequently, Esau was not concerned with the topographic differences between different bundles within the internode since these are largely controlled by the activity of the intercalary stem meristem, and she gave a detailed account of bundle development in leaves, which 1s not considered here. Despite this, similar features of development in the two plants include the method of early development of the procambium and the early appearance of protophloem and protoxylem at opposite poles of the strand (FiGureE 2). Increase in diameter of the bundle in both plants occurs by tan- gential longitudinal division that produces radial seriation of cells in transverse sections of the bundle (e.g., FiGure 2, D). Esau considered it inappropriate to view this as a ‘““cambium” (an opinion with which we concur) since this implies homologies with vascular organization that are not likely to exist in dicotyle- dons. That this is not a cambium is clear from the observation that division occurs during elongation of the vascular bundle. FiGure 2, D, for example, in which radial seriation is particularly obvious, represents a level only 1 mm below the shoot apex, whereas extension occurs over an additional distance of several centimeters (cf. FiGuREs | and Esau indicated that at any one level, proreonioenl always appears first, pro- 1984] TOMLINSON & VINCENT, RHAPIS EXCELSA, X rae | toxylem second. This sequence is common in stem bundles in Rhapis, but it is not always followed due to the greater topographic complexity. Protoxylem may precede protophloem (FiGure 2, C) in the differentiation of leaf traces, and metaxylem initiation in axial bundles may even precede initiation of proto- phloem. Metaxylem initials, of course, are not totally mature until much later, after axial elongation 1s complete Despite these differences, some general features of vascular differentiation in Rhapis—including the continuous acropetal differentiation of phloem and metaxylem—correspond to those found in other angiosperms. In contrast, pro- toxylem is discontinuously differentiated within each bundle, originating in the distal part of each leaf trace and, in a relative sense, rapidly extending both acropetally and basipetally. The protoxylem of the proximal leaf trace must advance acropetally in the stem to make contact with the distal protoxylem of the bundle in the leaf base. Although it is reasonable to assume topographic advance of further protoxylem differentiation in a basipetal direction, we have not estimated this quantitatively against fixed reference points. There is never continuity between protoxylem of one trace and that of another. This discon- tinuity is a topographic consequence of the acropetal advance of the cessation of internodal elongation and a direct expression of stem growth via an unin- terrupted meristem. Continuity of the xylem transport pathway is effected by the juxtaposition of metaxylem and the last-formed protoxylem. This is the basis for the ‘thydraulic constriction“ at each leaf insertion, recognized by Zimmermann and Sperry (1983) and illustrated in Zimmermann et al. (1982), upon which the whole hydraulic architecture of the palm is based. This simple developmental arrangement is reinforced by the failure of metaxylem to dif- ferentiate in the outgoing leaf trace beyond the level of departure of the last bridge or axial bundle. PHLOEM DIFFERENTIATION Unlike Esau, in her study of Zea (1943), we have been unable to make a very clear distinction between protophloem and metaphloem. In Zea the latter is structurally distinct because only metaphloem includes companion cells and its elements are arranged in regular radial series. Such a distinction is not evident in Rhapis. However, if we accept that metaphloem, by definition, only matures after the bundle is completely elongated, it is clear from FiGure 5 that all the phloem differentiated in the crown is protophloem. Obliteration of first- formed protophloem iS extensive, but some passive extension can presumably be accommodated. Sinc and p hl are not topographically distinct (unlike en en and metaxylem), continuity of development is evident. LEAF TRACE AND AXIAL BUNDLE We have used the terms ‘“‘axial bundle’ and “‘leaf trace” without precise definition in this series of papers. “Leaf trace” refers to the outwardly curved portion ofa bundle shortly before its entry into the leaf base, and “axial bundle” to the portion of a bundle remote from its entry into a leaf. With progressive 212 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 increase in our understanding of the pieeeoae initiation, and differentiation of vascular bundles, the terms can now be refined somewhat. In a topographic sense, the axial bundle can be said to See a leaf trace either at the point of its maximum level of penetration into the stem center or at the level of departure of the continuing axial bundle. The latter is unsatisfactory because this varies considerably in different monocotyledons: the leaf trace would be very short in some because the axial bundle departs close to the stem periphery, and in others because a distinct axial bundle is not always present (Zimmer- mann & Tomlinson, 1974, fig. 9 In developmental terms the level at which the “‘basipetally” determined influence of the leaf conjoins the ‘“‘acropetally’’ determined influence of the meristematic cap could represent the junction of the two portions of a subse- quently continuous vascular bundle. This definition is difficult to apply, al- though it may have the most precise morphogenetic meaning. It corresponds closely to the first topographic definition. In histological and functional terms, the ‘‘leaf trace” can be defined precisely direct functional significance because it is the basis for the hydraulic constriction mentioned by Zimmermann and Sperry (1983). The definition differs from both preceding ones because the “axial bundle” (represented by the metaxylem) and the “‘leaf trace” (represented by the protoxylem) necessarily overlap con- siderably. The “axial bundle” is continuous along the stem and makes no direct contact with the leaf. We thus have a precise developmental and functional application of the abstract and much-debated notion of the ‘‘cauline bundle” (Esau, 1965). The “‘cauline’ portion corresponds to the axial vasculature in which metaxylem 1s differentiated, since this is the pathway for axial movement of water up the stem (FiGure 8, B). The “foliar” portion (FIGURE 8, A) is the leaf trace in the above protoxylic context and, at least in a palm, relates solely to the irrigation of the major appendages (the leaves) via the region of direct contact (FIGURE 8, C). Protoxylem discontinuity appears to be a universal developmental feature of stem vascular tissues (Esau, 1965) but has not previously been explained in a functional sense. Hydraulic architecture gives a possible clue. At times of stress, xylem dysfunction is restricted to disposable plant parts while axial continuity is preserved (cf. Zimmermann, 1983) From this analysis and its functional application, we can extract information that can be applied to vascular plants generally. For example, analyses of Populus show similarity in the time of appearance of vascular tissues and the development of hydraulic constrictions (Larson, 1976, 1982: Larson & Ise- brands, 1978). However, these features are much condensed in the apical region: events that occur over distances of a few hundred microns in Populus (Larson, 1975) extend over centimeters and throughout many plastochrones in palms. This is the basis for our suggestion that palms are particularly suitable organisms for studying vascular development. 1984] TOMLINSON & VINCENT, RHAPIS EXCELSA, X 213 ACKNOWLEDGMENTS Access to living plants at the Montgomery Foundation, Fairchild Tropical Garden, provided by John Popenoe and J. B. Fisher, is appreciated. Monika Mattmiiller and Martin Zimmermann are thanked for access to section series, films, and preliminary plots that they prepared. Frank Ewers and Martin Zim- mermann critically reviewed the manuscript; constructive reviews by P. R. Larson and P. M. Rury after submission of the manuscript are particularly appreciated. We also thank John Sperry for advice and field assistance in Miami. LITERATURE CITED Esau, K. ee Ontogeny of the vascular bundle in Zea mays. Hilgardia 15: 327-356. —. 1965. Vascular differentiation in plants. Holt, Rinehart and Winston, New ork. FisHer, J. B., & J. C. FReNcH. 1976. The occurrence of intercalary and uninterrupted meristems in the internodes of tropical monocotyledons. Amer. J. Bot. 63: 51 525 Hatté, F., R. A. A. OLDEMAN, & P. B. Tomiinson. 1978. Tropical trees and forests— an architectural — Springer Verlag, Berlin. Larson, P. R. 1975. velopment and organization of the primary vascular system in Populus anes a to phyllotaxy. Amer. J. Bot. 62: 1084-1099 Procambium vs. cambium and protoxylem vs. metaxylem in Populus deltoides seedlings. Ibid. 63: 1332-1348. 198 he concept of cambium. Pp. 85-121 in P. Baas, ed., New perspectives in ‘wood anatomy. Martinus Nijhoff/W. Junk, The Hague, Boston, and London .G. IseEBRANDS. 1978. Functional significance of the nodal constricted zone in Populus deltoides. Canad. J. Bot. 56: 801-804 PARTHASARATHY, M. V., & P. B. TOMLINSON. 1967. Observations on metaphloem in the ee parts ‘of palms. Amer. J. Bot. 55: 1140-11 Priest.ey, J. H., L. I. Scorr, & E. C. Grtterr. 1935. The development of the shoot in Alstroemeria and th t of shoot growth in nn. Bot. (London) 49: Peron Pp. B. 1984. Development of stem conducting tissues in monocotyledons. In: R. A. Waite & W. L. Dickison, eds., Contemporary problems in plant anatomy. Academic Press, New York (in press .H. Zimmermann. 1967. The “wood” of monocotyledons. Bull. Int. Assoc. Wood Anat. 1967(2): 4-24. ZIMMERMANN, M. H. 1983. Xylem structure and the ascent of sap. Springer Verlag, erlin. & M. Matrmuiccer. 1982. The vascular pattern in the stem of the palm Rhapis excelsa. 1. The mature stem. II. The growing stem. Films C1404 and D1418. Institut fiir den Wissenschaftlichen Film, Nonnensteig 72, 34 Gottingen, West Germany. K. F. McCue, & J. S. Sperry. 1982. An natomy of the palm Rhapis excelsa, VIII. Vessel network and vessel-length distribution in the stem. J. Arnold Arbor. 63: 83-95. &J.S. Sperry. 1983. Anatomy ofthe palm Rhapis excelsa, (X. Xylem structure of the leaf meristem. J. Arnold Arbor. 64: 599-609. 214 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 P. B. ToMuINson. 1965. Anatomy of the palm Rhapis excelsa, 1. Mature vegetative axis. J. Arnold Arbor. 46: 160-178. & 1966. Analysis of complex vascular systems in plants: optical shuttle method. Science 152: 72, 73. & ———. 1967. Anatomy of the palm Rhapis excelsa, 1V. Vascular develop- ment in apex of vegetative aerial axis and rhizome. J. Arnold Arbor. 48: 122-142. & 1972. The vascular system of monocotyledonous stems. Bot. Gaz. 133: 141- 155. & . 1974. Vascular patterns in palm stems: variation of the Rhapis principle. J. Arnold Arbor. 55: 402-424. HARVARD FOREST PETERSHAM, MASSACHUSETTS 01366 1984] ZARUCCHI, AUBLET’S GENERIC NAMES 219 THE TREATMENT OF AUBLET’S GENERIC NAMES BY HIS CONTEMPORARIES AND BY PRESENT-DAY TAXONOMISTS JAMES L. ZARUCCHI In 1775 Aublet published his [Histoire des Plantes de la Guiane Francoise, an illustrated account of his botanical studies in the French colony between 1762 and 1764. In this work Aublet described many new species and introduced 208 new genera of plants to science. In a recent work, Howard (1983) dealt with the identities of the plants illustrated in the 392 plates in the work, reviewed the various taxonomic problems or uncertainties concerning the iden- tities of some of Aublet’s species, and presented a selected list of critical ref- erences that consider Aublet or the plants that he collected and described. The purpose of the present paper is to review how several botanists who were contemporary with Aublet treated his 208 genera. Scopoli, in his /atro- ductio ad Historiam Naturalem (1777), and Schreber, in the two-volume eighth edition of Linnaeus’s Genera Plantarum (1789, 1791), proposed many new generic names as replacements for names of Aublet that they considered “vulgar or barbarous” (Stafleu, 1971). Although neither Scopoli nor Schreber treated all of Aublet’s genera, A. L. de Jussieu, another contemporary botanist, re- viewed all 208 generic names in his Genera Plantarum (1789) and placed nearly all of them within his system of natural families. Many additional replacement names for Aublet’s taxa were provided by Necker in his three-volume E/ementa Botanica (1790). Necker’s names are not treated in this paper since his “species naturalis’’ are considered to be monomial names for species and therefore not validly published (Stafleu & Cowan, 1981). The Sydney Code (Voss ef al., 1983) supports this view of the nomenclatural status of Necker’s names under Article 20.4(b). In addition to publishing the 208 generic names in his Histoire des Plantes de la Guiane Francoise, Aublet validated the pre-Linnaean name Clompanus of Rumphius, by describing a second species under that genus. The nomen- clatural and taxonomic problems associated with Clompanus [Rumph.] Aublet are presently being studied by Geesink (pers. comm.). Taste | presents Aublet’s 208 generic names in alphabetical order; lists their treatment according to Scopoli (1777), Schreber (1789, 1791), and Jussieu (1789); and gives their current disposition as presented by Howard (1983). Modifications or explanations are added where needed. Footnotes supply any additional nomenclatural or taxonomic information necessary TaBLe 2 provides an index by entry number to all of the generic names © President and Fellows of Harvard College, 1984. Journal of the Arnold Arboretum 65: 215-242. April, 1984. TABLE 1. The nomenclatural and taxonomic treatments of Aublet’s generic names by seas (1777), Jussieu (1789), and their current taxonomic statu Schreber (1789, 1791), and = Aublet. (1775) Scopoli (1777)* Schreber (1789, 1791)* Jussieu (1789)? Current Status” ‘ oo on NC NC Cananga Aublet Guatteria 5 aA. St. ("Abereno! in T (537J) . Ablania (ELAE) . Abuta (MENI) - Acioa (CHRY) i ioua" in T . +Acouroa (LEGU) =*Dalbergia L. f. [178TT) . Aiouea (LAUR) . Amaioua (RUBI) "on P) . Amanoa (EUPH) . Ambelania (APOC) . Aniba (LAUR) . t+Apalatoa (LEGU) ("Opalatoa" on P) (= *rudig Schreber [1789]) Ablania Aublet NC Acioa re (TAci in I) "Acovrea" Aublet ("Acourea in I) = Ehrhardia Scop. ~("Aiovea" in S) NC Amanoa Aublet = Benteca Adanson (1763) ("Ambel ami Ambelamia" in 1) Aniba Aublet + Waldschmidtia Scop. + Trichocarpus Schreber an GP CES Ga NC + Acia Schreber (1791) corr. in Addenda) NC + +Douglassia 8 Schreber (1791) NC NC + +Willughbeja 9 Schreber (1789, 1791) (with Pacouria Aublet) + Cedrota Schreber (1789) + Cyclas Schreber (1789) (with Touchiroa Aublet) Ablania Aublet Abuta Aublet Acioa Aublet Acouroa Auble (with vatairea Auble nS) "Ajovea" Aublet "Amatoua" Aublet Amanoa Aublet Ambelania Aublet Aniba Aublet Apalatoa Aublet (Touchtroa in S) § Ruiz & Pavon (1794) Sloanea L. (1753) Abuta Aublet® Acioa Aublet 2? *Dalbergia L. f. ¢ —_ 1781) Aiouea Aublet Amaioua Aublet Amanoa Aublet Ambelania Aublet Aniba Aublet *Crudia Schreber (1789) Ole WOLAYOdIAV ATIONAV AHL JO TYVNUNOL 69 “T0a] = as Mm ine) . Apeiba (TILI) . Arouna (LEGU) . Aruba (SIMA) 11 *Bacopa (SCRO) 2 . Bagassa (MORA) : Baer (COMP eria" in T 793 ) . Banara (FLAC) . Bassovia (SOLA) . Bertiera (RUBI) . Bocoa (LEGU) . Cabomba (CABO) . Cacoucia (COMB) + . Calinea (DILL) , +Cananga (ANNO) uatteria Ruiz a P avon [1794]) . Caraipa (BONN) ("Caraipe" in T [537]) . Carapa (MELT) Apeiba Aublet "Arovna" Aublet "Arvba" Aublet Bellonia L. (1753) NC "Bailliaria" ah ("Baillaria" in I) Banara Aublet Bassovia Aublet Bertiera Aublet NC Cabomba Aublet Hanbergera Scop. ("Cacoutia" in S) Calinea Aublet Cananga Aublet NC #Aubletia Schreber (1789) '° (with SToanea "Loefl.") NC NC Bacopa Aublet NC gi Sw. 1788) '2 ("Bailliera" in S) Banara Aublet Bassovia Aublet Bertiera Aublet NC Nectris Schreber (1789) NC Doliocarpus Rolander (1756 NC A eiba pee ia " Cait pe [17 ane Arouna Aublet Aruba Aublet Bacopa Aublet Bagassa Aublet "Ballieria" Aublet Banara Aublet Bassovia Aublet Bertiera Aublet Bocoa Aublet Cabomba Aublet Cacoucia Aublet Calinea Aublet Cananga Aubl et with Uvaria pongifolia conn: Aublet in § Caraipa Aublet (Caratpa in 1) Carapa Aublet® Apeiba Aublet Dialium L. (1767) Simaba Aublet *Bacopa Aublet Bagassa Aublet Clibadium L. (1771) Banara Aublet Solanum L. (1753) Bertiera Aublet Bocoa Aublet Cabomba Aublet *Combretum Loefl. (1758) Doliocarpus Rolander (1756) *Guatteria Ruiz & Pavon (1794) Caraipa Aublet'? Carapa Aublet [p86l SHNVN OLANAD S.LATANV ‘THOONUVZ LIG TABLE | (continued). Aublet (1775 ) Scopoli (1777)* Schreber (1789, 1791) 2 Jussieu (1789)? Current Status 4 . +Carapichea (RUBI) (=*Cephablis Sw. [1788 y . Cassipourea (RHIZ) . Catinga (MYRT) ("Catringa" - Ciponima (SYM in 1) P) CS Siponima" on P) . +Coublandia (=*Mueller [1 . Couma (APOC ) . Cipura (IRID) . Conami (EUPH) ( a : eonceveiba (EUPH) ( ao _ in T > &on . Conobea (SCRO) .(+)Conohoria (VIOL) LEGU) . f. . Couepia (CHRY) P) 16 2 * a = Chesnea Scop. Tita Scop. NC "Siponima" Aublet "Cipvra" Aublet NC NC Conobea Aublet Conohoria Aublet ("Conohria" in = ?Caesalpinia L. 1) (1753) + Callicocca Schreber Hf a a + (wi & Ever Au ae [*Ce are Sw 1788) 3 adde ss in the "addenda ied eee: (1791) - a S h Tapogomea Aub] Legnotis Sw. (1788) '4 NC Sym socos Seg (1760 Sipon in S) Marica Schreber (1789) NC NC Conopea Schreber (1791) "Conolea" in I) NC NC Acia Schreber (1791) Conepia" rein kde nda )’ ) = Tapogomea Aublet = Cassipourea Aublet Catinga Aublet Ciponima Aublet Cipura Aublet Conami Aublet® § Conceveiba Aublet Conobea Aublet Conoria A. L. Juss. Coublandia Aublet Couepia Aublet Couma Aublet® *Psychotria L. (1759) Cassipourea Aublet ? Calycorectes SS Be . Berg Symplocos Jacq. Cipura Aublet Phyllanthus L. (1856) (1760) 1753) Conceveiba Aublet Conobea Aublet (*)Rinorea Aublet '® *Muellera L. f. Couepia Aublet Couma Aublet (1781) 15 8IC WOLAYOPUV ATIONYV AHL JO TWNUNOfL $9 “10A] > nn on WM On oO . +Coumarouna (LEGU) + Heinzia Scop. + *Dipteryx Schreber (1791) Coumarouna Aublet *Dipteryx Schreber (1791) (=*Dipteryx (with Taralea Aublet) Schreber [1791]) .(+)Coupoui (ruBr) | NC NC Coupoui Aublet *Duroia L. f. (1781) 17 . Couratari (LECY) NC NC = Zanonia L. (1753) Couratari Aublet . Courimari (?ELAE) NC NC Courimari Aublet ? Sloanea L. (1753) 8 . Couroupita (LECY) * Poon. eana Scop. NC Couroupita Aublet Couroupita Aublet ("Coroupita" in S) . Coussapoa (CECR) NC NC Coussapoa Aublet Coussapoa Aublet . Coussarea (RUBI) + Pecheya Scop. NC Coussarea Aublet Coussarea Aublet . Coutarea (RUBI) "Covtarea" Aublet NC Coutarea Aublet Coutarea Aublet ( h Portlandia "Jacq. L. . B [1756] in S) . Coutoubea (GENT) + Limborchia Scop. + Picrium Schreber (1791) Coutoubea Aublet Coutoubea Aublet . Crenea (LYTH) "Crena" Aublet "Crenaea" Aublet Crenea Aublet Crenea Aublet (Cranea" in I) . +Deguelia (LEGU) "Degvelia" Aublet NC Deguelia Aublet *Derris Lour. (1790) =*Derris Lour. £1790J) . Enourea (SAPI) "Enovrea" Aublet NC Enourea Aublet Paullinia L. (1753) . Eperua (LEGU) "Eperva' Aublet + Dimorpha Schreber (1791) Eperua Aublet Eperua Aublet Tam Par Woe Aublet) . +Evea (RUBI) Evea Aublet + Callicocca Schreber (1789) Evea Aublet Faramea Aublet =*Cephaélis Sw. (with Tapogomea Aublet [1788 & Carapichea Aublet) . Faramea (RUBI) = Knoxia L. (1753) NC Faramea Aublet Faramea Aublet . +Ferolia (MORA) NC NC = Parinarium A, L. Juss. *Brosimum Sw. (1788) (=*Brosimum Sw. (see under no. 117) 88]) [p86 SAWVN OWA NAD S.LATANV IHOONUVZ 61C TABLE | (continued). fon) Aublet (1775) Scopoli (1777)¢ Schreber (1789, 1791)* Jussieu (1789)° Current Status“ 56. #Fothergilla (MELA) '? + +Leonicenia Scop.22 * Melastona L. (1753) = Melastoma “Burm. L." *Miconia Ruiz & Pavon mu arious genera (1794) 57. Galipea (RUTA) NC NC "Galipaea" Aublet Galipea Aublet 58. Goupia (GOUP) "Govpia" Aublet + Glossopetalum Schreber Goupia Aublet Goupia Aublet ("Guopia" in 1) C785} 59. Guapira (NYCT) "Gvapira" Aublet _NE = Avicennia L. (1753) Pisonia L. (1753) ("Quapira" on P) 60. Hevea (EUPH) NC + Siphonia D. Rich. ex = Jatropha L. (1753) Hevea Aublet Sc hreber (1791) . Heymassoli (OLAC) + +Rottboelia Scop.2! NC = Ximenia "Pl. L." (1753) Ximenia L. (1753) 62. +Houmiri (HUMI) + Wernisekia Scop. + Myrodendrum Schreber + Houmiria A. L. Juss. “or Aublet corr. (=*Humiria Aublet (1789) . ot. Hil. (1805) corr. J. - Hil. [1805]) ("Homiri" in T [531]) 63. +Icacorea (MYRS) NC NC Icacorea AubletS *Ardisia Sw. (1788)¢2 (=*Ardisia Sw. (17887) . Icica (BURS) Icica Aublet NC Icica Aublet Protium Burm. f. (1768) 65. Iroucana (FLAC) + Moelleria Scop. + #Athena 33 = Anavinga "“Rheed. Lam." Casearia Jacq. (1760) Schrebes (1789) . Ivira (STER) Ivira Aublet = Sterculia L. (1753) = Sterculia L. (1753) Sterculia L. (1753) ("Tuira on P) . Licania (CHRY) Licania Aublet + Hedycrea Schreber (1789) Licania Aublet Licania Aublet 68. Licaria (LAUR) NC NC Licaria Aublet Licaria Aublet . Mabea (EUPH) Mabea Aublet Mabea Aublet Mabea Aublet Mabea Aublet WOLAYOUdV WIONAV FHL JO TVWNYNOL OCC ¢9 “TOA] —s —S ~s Sw Frans | oy . Macahanea (?CELA) NC NC + Macanea A. L. Juss. ? Salacia L. (1771) . Macoubea (APOC) NC NC Macoubea Aublet Macoubea Aublet . Macoucoua (AQUI)** + 4Labatia Scop.” NC = Ilex L. (1753) lex L. (1753) racoua" in T Macou yeaa “Auble t in 3]) ucava" in I) . Mahurea (BONN) "Mahvrea" Aublet 7 ee Schreber (1789)*° Mahurea Aublet Mahurea Aublet ee hur a" Aublet 1) . Maieta (MELA) Maieta Aublet = Melastoma L. (with "Mayeta" Aublet Maieta Aublet ayeta" in T various genera in S) on P) ("Mayeta" Aublet in T & I) . Malanea (RUBT) = Chomelia L. (1758) + +Cunninghamia 27 Malanea Aublet Malanea Aublet Schreber (1791) TESS in I; with "Antirrhea nee. a) . Manabea (VERB) = Pavetta L. (1753) [RUBI] = Aegiphila Jacq. (1767) Manabea Aublet Aegiphila Jacq. (1767) . Managa (?CELA) NC NC Managa Aublet ? Salacia L. (1771) . Mapania (CYPE) Mapania Aublet NC Mapania Aublet Mapania Aublet - Mapouria (RUBI) “Mapovria" Aublet NC = Simira Aublet *Psychotria L. (1759) . Maprounea (EUPH) NC = Aegopricum L. (1775) Maprounea Aublet Maprounea Aublet ("Mapronnea" on P) . Maquira (MORA) NC NC Maquira Aublet® Maquira Aublet . Maripa (CONV) NC NC Maripa Aublet Maripa Aublet : Matayba Ahi + Ernstingia Scop. + Ephielis Schreber ee Matayba Aublet Matayba Aublet ("Mataiba" on P) Mataiba" in T & I) ("Mataiba" in I) . Matelea (ASCL) Matelea Aublet NC Matelea Aublet Matelea Aublet . Matourea (SCRO) "Matovrea" Aublet = Vandellia P. Br. [ex L.] Matourea Aublet *Stemodia L. (1759) gen. 706) (1767) + Dickia Scop. (gen. 877) SHNVN OIYANAD S.LATANV THOONUVZ [p861 GG I TABLE | (continued). Aublet (1775) Scopoli (1777)* Schreber (1789, 1791)¢ Current Status? @ a wo fon) . Mayaca (MAYA) . +Mayepea (OLEA) + ("Mayepa" i . Mayna (FLAC) . Meborea (EUPH) . Minquartia (OLAC) . Montira (LOGA) . Moquilea (CHRY) = . Moronobea (GUTT) im . Mourera (PODO) . Mouriri (MELA) + ‘. Mommoucoa (CONV) 7 ("Mouroucoua" in 1) . Moutabea (POLY) ("“Moutoubea" in I) Mayaca Aublet Freyeria Scop. ("Mayepa" Aublet in S) NC Meborea Aublet NC Lea Aublet ("Montiva" in 1) "Dahvronia" Scop. Oguilea" in S) Blakstonia Scop.°? "Movrera" Aublet Bockia Scop. (ovriri" in S) Maireria Scop. NC = — + + + a r Symphonia L. f. Syena Schreber (1789) +Ceranthus Schreber (1789 NC Rhopium Schreber (1791) 1781) Lacis Schreber (1789) Myrtus L. (1753) “(with v arious genera in S) (gen. 844 in 1789) = — Sw. (1788) (ge 750 in 1791) NC + Cryptostomum Schreber (1789) Mayaca Aublet Chionanthus L. (1753)28 Mayna Aublet ? Phyllanthus L. (1753) Minquartia Aublet Spigelia L. (1753) Licania Aublet Moronobea Aublet Mourera Aublet Mouriri Aublet Maripa Aublet Moutabea Aublet ce WOLAYOUUV AIONYV AHL JO TVNANOL 69 “10A] wo wo . Moutouchi (LEGU) oe Scop. NC = +Pterocarpus arte L 130 *Pterocarpus Jacq. (1763) ( ‘Mourouchi uchi" in I) ("Moutouchiam' "Moutouchi a" . Nacibea (RUBI) Nacibea Aublet = *Manettia Mutis [ex L.] Nacibea Aublet *Manettia Mutis ex L. (also cited +Lygistum (*Man nctra "Mut. Lb." (771) P. Br. C1756] in S)_ C71 S) . Napimoga (FLAC) + Tattia Scop. NC Napimoga Aublet Homalium Jacq. (1760) . Nonatelia (RUBI) Nonatelia Aublet + Oribasia Schreber (1789) Nonatelia Aublet *Psychotria L. (1759) . Norantea (MARC) Norantea Aublet + Ascium Schreber (1789) Norantea Aublet Norantea Aublet . Ocotea (LAUR) Ocotea Aublet = Ponastet cas (1791) Ocotea Aublet Ocotea Aublet iE tandra ers [ex Rot ott 11778] ) . #0relia (APOC) = Gentiana L. 53) = Allamanda L. (1771) = Allamanda L. (1771) Allamanda L. (1771) illeg. name oe ee ee ‘donson (with "G balarips a on All amanda and Swertia L.) D. Allamand.” [ex L.]) [177T]) . *Ouratea (ocHn)>! "Ovratea" Aublet NC Ouratea Aublet *Quratea Aublet : ey = Genipa L. (1754) + *Uncaria Schreber = Nauclea L. (1762) *Uncaria Schreber (1789) (=*Unc oe ae (1789) C7851) . +Outea GU) "Ovtea" Aublet + *Macrolobium Schreber Outea Aublet *Macrolobium Schreber =*M crolcbiun (1789) Schreber [1789]) (with Vouapa Aublet) . Pachira (BOMB) Pachira Aublet = Carolinea L. [f.] (1781)22 Pachira Aublet?? Pachira Aublet . +Pacouria (APOC) + +Alstonia Scop." - emi aghbeda Schreber?” Pacouria Aublet +Pacouria Aublet (=*Landolphia (1789, corr. 1791) P. Beauv. [1805]) (with MAbenaGi Aublet) . Pacourina (COMP) + Meisteria Scop. NC Pacourina Aublet Pacourina Aublet . Pagamea (R NC NC Pagamea Aublet Pagamea Aublet UBI) "Pagama" in T a corr. in Addenda) [P3861 SHNVN OFWANAD S.LATANV ‘THOONUVZ EGC TABLE | (continued). Aublet (1775) Scopoli (1777)? Schreber (1789, 1791) 2 Jussieu (1789)? Current Status” = Ie . Palicourea (RUBI) . Paloue (LEGU) ("Palove" in I) . [+]Pamea (COMB) * . Paralea (EBEN) . Pariana (GRAM) . Parinari (CHRY) . Parivoa (LEGU) . Passoura (VIOL) . Patabea (RUBI) . Patima (RUBI) . Paypayrola (VIOL) . Pekea (CARY) . Perama (RUBI) . Perebea (MORA) = Simira Aublet urea" in S, + Ginannia Scop. NC NC Pariana Aublet "Dvgortia" Scop. Parivoa Aublet NC +"Sicelivm" P. Br 2 i (with Tontanea Aublet) NC + Lignonia Scop. Pekea Aublet NC ge Aublet ("Peribea" in I) Stephanium Schreber (1789) {" ae in corr. in Addenda a 1791]) Ginannia Scop. (1777) ("Palouk" Aublet in Trt NC NC Pariana Aublet + Petrocarya Schreber (1789) + Dimorpha Schreber (1791) Ta or a Aublet) + Rhizobolus Gaertner ex Schreber (1789) + Mattuschkaea Schreber 1791 NC get Aublet (w Failed Aublet) + Palovea (w ee «Paloue” heBIe Pamea Aublet Paralea Aublet Pariana Aublet = Parinariu “Ww ith Ferolia eo , Parivoa Aublet = Riana Aublet Patabea Aublet Patima Aublet + Payrola A. L. Juss. nekes Aublet ms ae Sr Aublet Perama Aublet Perebea Aublet Palicourea Aublet Paloue Aublet Buchenavia 36 Fichler (1866 ) Diospyros L. (1753) Pariana Aublet Parinari Aublet Eperua Aublet Rinorea Aublet Ixora L. (1753) Patima Aublet Paypayrola Aublet Caryocar L. (1771) Perama Aublet Perebea Aublet pec WOLAYOUUV GATONYYV AHL JO TYWNANOFL $9 “10A] = ine) Lon) . #Pirigara (LECY) . Posoqueria (RUBI . Pouteria . Piparea (FLAC) . a (MORA ) 1788 um Sw. 38 . Piripea (SCRO) . Piriqueta (TURN) . Pitumba (FLAC) . Poraqueiba (ICAC) : — (LEGU) artzia Schreber . Potalia (LOGA) . Pourouma (CECR) (SAPO) - Qualea (VOCH) . Quapoya (GUTT) . Quararibea (BOMB) ~ * * + i a + a Pirigara Aublet Piripea Aublet "Byrghartia" Scop. ("Pariqueta" in S) NC et Scop 40 ( Paraaieiba” in S) Ramepekia Sc cop. Posogueria" Possira Aublet in S) Potalia Aublet NC Pouteria Aublet "Qvalea" Aublet eae 44 +Smithia Scop. ("Gvapoya" in S) yeclles Scop. raibea" in S; iqusvaiba in 1) 2 * Buchnera L. *Gustavia L. (1775) (1753) "Burcardia" Scop. +Wolfia Schreber (1791) #Barreria Scop. Cyrtanthus Schreber (1789) Rittera Schreber (1789) #Nicandra Schreber (1789)" NC *haetocarrs Schreber (gen. 179 in 1789) *Labatia Sy. (1788)°3 (gen. 1724 in 1791) Qualea Aublet Xanthe Schreber (1791) 46 Myrodia Sw. (1788) 39 aay Piparea Aublet Piratinera Aublet § Pirigara Aublet mc th Gustavia "LS." nS) Piripea Aublet Piriqueta Aublet Pitumba Aublet® Poraqueiba Aublet ("Poraqueiba" in Posoqueria Aublet m Possira Aublet | potalia Aublet Pourouma Aublet Pouteria Aublet Qualea Aublet Quapoya Aublet Quararibea Aublet Casearia Jacq. (1760) *Brosimum Sw. (1788) 38 *Gustavia L. (1775) Buchnera L. (1753) Piriqueta Aublet Casearia Jacq. (1760) Poraqueiba Aublet Posoqueria Aublet *Swartzia Schreber (1791) Potalia Aublet Pourouma Aublet Pouteria Aublet Qualea Aublet Quapoya Aublet Quararibea Aublet [p86 SHINVN OINANAD S.LATANV THOONUVZ SCC TABLE | (continued). Aublet (1775) Scopoli (1777)? Schreber (1789, 1791) Jussieu (1789)? Current Status® . Quebitea (PIPE) NC NC = Dracontium L. (1753) Piper L. (1753) ARAC - Quiina (QUIT) NC NC Quiina Aublet® Quiina Aublet . Racaria (SAPI) NC NC Racaria Aublet Talisia Aublet . Racoubea (FLAC) + "Lagvnezia" Scop. = Homalium Jacq. (1760) = Homalium Jacq. (1760) Homalium Jacq. (1760) . Rapanea (MYRS) = Plotia Adanson (1763) NC Rapanea Aublet Myrsine L. (1753) . Rapatea (RAPA) Rapatea Aublet + Mnasium Schreber (1789) Rapatea Aublet Rapatea Aublet . Raputia (RUTA) "Rapvtia" Aublet + Sciuris Schreber (1789) Raputia Aublet Raputia Aublet . [+]Remirea (cypE)*” "Remeira' Aublet + Miegia Schreber (1791) Remirea Aublet *Mariscus Vahl (1805/6) 47 . Riana (VIOL) = Rinorea Aublet NC Riana Aublet Rinorea Aublet (with Passoura Aublet in S) sana 48 , ; ; 48 . (*)Rinorea (VIOL) Rinorea Aublet NC Rinorea Aublet (*)Rinorea Aublet (with Riana Aublet in S) 1. Ronabea (RUBI) "Ronobea" Aublet NC Ronabea Aublet *Psychotria L. (1759) onobea" in I & on P) . Ropourea (EBEN) + +Idesia Seon. + Camax Schreber (1789) Ropourea Aublet Diospyros L. (1753) . Rouhamon (LOGA) + Chemnicia Scop. + Lasiostoma Schreber = Strychnos L. (1753) Strychnos L. (1753) ("Rohamon" in S) (1789) (with Ignatia "LS." 'Caniram Rheed." in S) . Roupala (PROT) + Leinkeria Scop. "Rhopala" Aublet Roupala Aublet Roupala Aublet . *Rourea (conn)?! "Rovrea" Aublet + Robergia Schreber (1789) Rourea Aublet *Rourea Aublet WOLIYOPUUV GATIONUYV AHL AO TVNANOL 977 $9 “10A] a . Sabicea (RUBI) . Sagonea (HYDR) . Saouari (CARY) . Senapea (?PASS) . Simaba (SIMA) : 53 . *Simarouba (SIMA) . Simira (RUBI) - Singana (?LEGU) . Sipanea (RUBI) . Siparuna (MONI) . Soramia (DILL) ia" in T . Souroubea (MARC) . Tachia (GENT) . Tachibota (CHRY) . Tachigali (LEGU) a * << aa Hamelia Jacq. Sagonea Aublet NC NC Simaba Aublet NC Simira Aublet Singana Aublet Sipanea Aublet NC Soramia Aublet +Loghania Scop. Tachia Aublet NC "Cvba" Scop. = a Schwenkfelda Schreber (17835) (gen. 306) Schwan teldie rego (179T) (co n Addenda) Reichelia Schreber (1789) in S) ("Sagona" i oa L. (1762) (author as as, ") NC NC #Zwingera Schreber (1791) NC NC Sterbeckia Schreber (1789) Virecta L. (1775)°" NC Doliocarpus Ro (1756) (gen. 874 in 1789) ander #Mappia Schreber?” gen. 1755 in 1791) Ruyschia Jacq. (1760) Myrmecia Schreber (1789 Salmasia Schreber (1789 "Cubaea" Scop. (1777) ~~ Sabicea Aublet Sagonea Aublet Pekea Aublet Senapea Aublet) Simaba Aublet Quassia L. (1762) Smira Aublet th Mapouria Aublet & Pal icone: Aublet Singana Aublet Mussaenda L. (a7e3i°° Siparuna Aublet Soramia Aublet Souroubea Aublet Tachia Aublet Tachibota Aublet "Tachigalia" Aublet Sabicea Aublet *Hydrolea L. (1762) Caryocar L. (1771) ? Senapea Aublet Simaba Aublet Simarouba Aublet Simira Aublet ? Singana Aublet Sipanea Aublet Siparuna Aublet Doliocarpus Rolander (1756) Souroubea Aublet Tachia Aublet Hirtella L. (1753) Tachigali Aublet?© [p86 SHAWN OFNANAD S.LATENV THOONUVZ LOG TABLE | (continued). 8CC Aublet (1775) Scopoli (1777) Schreber (1789, 1791)* Jussieu (1789)° Current Status” 171. +Taligalea (VERB) Taligalea Aublet = *Amasonia L. [f.] (1781) Taligalea Aublet *Amasonia L. f. (1781) masonia f [1784 172. Talisia (SAPI) Talisia Aublet NC Talisia Aublet Talisia Aublet 173. Tamonea (VERB)>” Tamonea Aublet + Ghinia Schreber (1789) Tamonea Aublet Tamonea Aublet?? (with "Verbena L." £1753] in S) 174. Tampoa (CELA) NC NC Tampoa Aublet’ ? Tampoa Aublet 175. Tanibouca (COMB) NC NC Tanibouca Aublet Terminalia L. (1767) 176. +Taonabo (THEA) + "Dvpinia" Scop. = *Ternstroemia Mutis ex + Tonabea A. L. Juss. *Ternstroemia Mutis ex =*Ternstroemia L. [f. 1781) L. f. (1781) Mutis ex L. f. a additional genera (17811). S) 177. Tapirira (ANAC) "Taparira" Aublet + Joncquetia Schreber (1789) "Tapiria' Aublet Tapirira Aublet 178. +Tapogomea (RUBI ) Tapogomea Aublet + call icocca Schreber (1789) Tapogomea Aublet *Psychotria L. (1759) (=*CephaEli (with Carapichea Aublet ith Carapichea Aublet (1 Aublet) S) 179. Tapura (DICH) "Tapvra" Aublet + Rohria Schreber (1789) Tapura Aublet Tapura Aublet 180. +Taralea (LEGU) Taralea Aublet + *Dipteryx Schreber (1791) Taralea Aublet *Dipteryx Schreber (1791) ipteryx Schreber (with Coumarouna Aublet) (1 94 181. +Tariri (SIMA) NC NC = Comocladia "Brown. L. *Picramnia Sw. (1788) (=*Picramnia Sw. Jacq P. Br.] (1756) 17887) Rte Pseudo-Brasiliu ne i Adanson (1763) ] 182. Thoa (GNET) Thoa Aublet Thoa Aublet Thoa Aublet Gnetum L. (1767) WOLAYOIUV ATONYY JHL JO TWNANOL $9 10a] @ . Tibouchina (MELA) + Savastania Scop. = Melastoma L. (1753) Tibouchina Aublet Tibouchina Aublet ("Tibochina" in 1) (with various genera in S) . Ticorea (RUTA) Ticorea Aublet + Ozophyllum Schreber (1791) Ticorea Aublet Ticorea Aublet . Tigarea (DILL) "Tigvrea" Aublet + Rhinium Schreber (1791) Tigarea Aublet Tetracera L. (1753) . Tococa (MELA) Tococa Aublet = Melastoma L. (1753) Tococa Aublet Tococa Aublet “with verou genera . Tocoyena (RUBI) ?= Genipa L. (1754)° NC Tocoyena Aublet Tocoyena Aublet . Tonina (ERIO) Tonina Aublet + Hyphydra Schreber (1791) Tonina Aublet Tonina Aublet . Tontanea (RUBI) = +"Sicelivm" P. Br. + #Bellardia 62 Tontanea Aublet *Coccocy selun P. Br 1 Schreber (1791) (1756 Schreber (with Patabea Aublet) (1791) . Tontelea (CELA) "Tontalea" Aublet + Tonsella Schreber (1789) Tontelea Aublet Tontelea Aublet . Topobea (MELA) Topobea Aublet NC Topobea Aublet Topobea Aublet . +Touchiroa (LEGU) NC : cyclas Schreber (1789) = Apalatoa Aublet *Crudia Schreber (1789) (=*Crudia Schreber ith Apalatoa Aublet) [1789]) - Toulicia (SAPI) "Tovlicia" Aublet + Ponaea Schreber (1789) Toulicia Aublet Toulicia Aublet ("Toulipita" in T ; a (LEGU ) NC + *Swartzia Schreber (1791) Tounatea Aublet *Swartzia Schreber (1791) F maizia tzia Schreber 63 . Touroulia (QUIT) + +Robinsonia Scop. = +Robinsonia Scop. (1777) Touroulia Aublet Touroulia Aublet . Tovomita (GUTT) NC NC Tovomita Aublet Tovomita Aublet . Trigonia (TRIG) Trigonia Aublet Trigonia Aublet Trigonia Aublet Trigonia Aublet . Vantanea (HUMI) NC + Lemniscia Schreber (1789) Vantanea Aublet Vantanea Aublet [p86 SAINVN OFYANAD S.LATANV THOONUVZ 6CC TABLE | (continued). 2 Aublet (1775) Scopoli (1777) Schreber (1789, 1791)¢ Jussieu (1789)°9 Current Status’ . Virola (MYRI) . Votomita (MELA) . Voyria (GENT) . Xiphidium (HAEM) . Vatairea se ) NC ("Voitai Virola Aublet . +Vochy (Voch) + Salmonia Scop. (=*Voch me Aublet ("Vochi" in T; "Vochy" in 1) corr. Poiret a a = [nom. & orth S:.1) Votomita Aublet . +Vouacapoua (LEGU) NC (=*Andira A. L. Juss. [1789]) . +Vouapa (LEGU) + "Krvegeria" Scop. (=*Macrolobium Schreber [1789]) . Vouarana (SAPI) NC . Voyara (CAPP) NC Voyria Aublet "Xiphidivm" Aublet NC 7 eee Thunb. ica Gronov. me + Cucullaria Schreber (1789) + Glossoma Schreber (1791) NC + *Macrolobium Schreber 1789 (with Outea Aublet) NC NC + Lita Schreber (1791) Xiphidium "Loefling. Tbl. + + Acouroa Aublet Virola Aublet Vochisia A. L. Juss. Votomita Aublet Andira ae ages . Lam. ex A. L. Juss.] (' ingelin Pl." in 5) Vouapa Aublet Ornitrophe Comm. ex A. L. Juss. ] ("Voua-rana" Aublet in $) Voyara Aublet "Vohiria" Aublet Xiphidium "Loefl. Aubl." Vatairea Aublet Virola Aublet *Vochysia Aublet corr. Poiret (1808) Votomita Aublet +Vouacapoua Aublet *Macrolobium Schreber (1789) Vouarana Aublet Capparis L. (1753) Voyria Aublet Xiphidium Aublet WOLAYOIUV ATONAYV FHL JO TWNUYNOL OE? C9 “10A] ‘This Table lists Aublet's 208 generic names in alphabetical order, followed by a four-letter le tea for the family (see Table 3). Conserved genera are preceded by an asterisk (*), rejected names by a plus sign (+), superfluous names by "+," and later homonyms by "4" In the case of a rejected name, its conserved counterpart is listed ee that entry in the first ein Orthographic variants apyeet in co followed by their location in the relevant publication: text (T), index (1), or plate (P). Names appearing in synonymy re cited a MSs, escopol i and Schreber did not consider all of Aublet's genera in their treatments; the genera that were not cited (NC) are so noted. 3the names with a section symbol (§) indicate the genera listed by Jussieu (p. 446) under "OBS. Praeter genera ordinis indeterminati." 4 according to the So provided by Howard (1983), except where noted. A question mark indicates doubt as to the identity or placement of a particular taxo oA Although Howard (1983) gives the identification as Guatteria, he mentions the possibility that Aberemoa may actually represent a species of Duguetia, against which Aberemoa is rejected. Osee notes on the lectotypification of Abuta by Barneby & Krukoff (1971) and by Howard (1983). “Howard (1983) indicates that there is no modern placement of Acouroa Aublet; the genus is rejected in favor of Dalbergia L. f. 8Not +Douglassia Miller (1754) -- Verbenaceae. Ischr eber attributed the authorship of +Willughbeja (not *Willughbeia ar [1820] -- Apocynaceae) to Scopoli in Volume 1 (1789), but later eee himself as the sole author in the Addenda found in Volume 2 (1791). not Aubletia Gaertner (1788) -- Sonneratiaceae. Ip acopa is conserved against +Moniera P. Br. (1756) and +Brami Adanson (1763) -- Scrophulariaceae. VeNot Trixis P. Br. (1756) -- Compositae; not #Trixis Adanson (1763) -- Haloragaceae. e four species of C Caraipa described by Aublet, only the Pars species, vifolia Aublet, is a member of the Bonnetiaceae. The eee species represent members of Licania and Couepia (chryaoEgT eeeaeT see Howard (1983) and Kubitzki (1978) for further details "4 agnotis was proposed by Swartz (1788) as a replacement name for Cassipourea Aublet. ferences relating to the as of Catinga (with lectotype species C. moschata Aublet), see Howard (1983). If Catinga Aublet and Calycorectes 0. Berg pro ihe be congeneric, then a choice must be made either to conserve the latter name or to make the appropriate combinations under Catinga SAINVN OFYdNaD S.LATANV THOONUVZ [p86 [eG 16 Hekking (1982) proposes that Rinorea Aublet be conserved over Conohoria Aublet. 17 A proposal to conserve Duroia L. f. over Coupoui Aublet will be submitted to Taxon (Zarucchi, in prep.). 18 rte while Aublet's work was being printed. ork were replaced after the initial page impress The identity of Courimari Aublet as a species of Sloanea L. is highly questionable; see Howard (1983) for further comments. Fothergilla L. (1774) -- Hamamelidaceae. Howard (1983) provides an account of the change from Tamonea (Melastomataceae) to a ae anes of various sano a Aublet (1775) has shown that several pages in 20 Leonicenia Scop. is rejected in favor of *Miconia Ruiz & Pavon (1794) -- Melastomataceae. 21 2 In combinations in Icac Not *Rottboellia L. f. (1780) -- Gramineae. a recent a Lundell —~ 1981) accepts Icacorea Aublet as distinct from Ardisia Sw. and published approximately one hundred 23 Not +Athenaea Adanson (1763) -- Compositae; not *Athenaea Sendtner (1846) -- Solanaceae. 24 In Index Nominum Genericorum, Farr and colleagues (1979) list the family as Celastraceae. Not *Labatia Sw. (1788) -- Sapotaceae. Not *Bonnetia Martius (1826) -- Bonnetiaceae. Not *Cunninghamia R. Br. (1826) -- Taxodiaceae. For details regarding the taxonomic standing of Mayepea Aublet, Linociera Sw. ex Schreber, and Chionanthus L., see Stearn (1976). Not Blackstonia Hudson (1762) -- Gentianaceae. Not *Pterocarpus Jacq. (1763) -- Leguminosae. . Ouratea Aublet is conserved, but no genus is listed as rejected in favor of Ouratea; see Voss et al. (1983). WOLAYOUUV CTIONYV AHL AO TVNUNOFL CEC $9 “10A] 32 carolinea L. f. is an illegitimate name; it was superfluous when published. 33carolinea "L.S." is listed as a replacement generic name. 34Not *Alstonia R. Br. (1810) -- Apocynaceae. Sap notes regarding this replacement name under Ambelania (entry no. 9). . a Aublet and Buchenavia Eichler are indeed congeneric, then either Buchenavia should be proposed for conservation or the ‘apeace ae ii under Pamea should be made ——— Siceliu P. Br. is rejected in favor of *Coccocypselum P. Br. (1756) corr. Schreber (1791). illegitimate when published since Japarandiba (a Marcgrave name validly ae aie by Adanson in 1763) was listed in rigara was is conserved against Japarandiba Adan Peecieiley Pirigara tetrapetala Aublet; Gustavia L. 39Not *Wolffia Horkel ex Schleiden (1844) -- Lemnaceae. Not Barreria L. (1753) -- Rutaceae. Not *Nicandra Adanson (1763) -- Solanaceae. MeNot *Chaetocarpus Thwaites (1854) -- Euphorbiaceae. 43 see entry no. 72). Not +Labatia Scop. (1777), a replacement name for Macoucoua Aublet (Aquifoliaceae) ( “4Not *Smithia Aiton (1784) -- Leguminosae. “SNot *Gerbera L. (1758) -- Compositae. Mcvartz (1788) introduced Myrodia as a replacement name for Quararibea Aublet. My How (1983) reports that the monotypic femjrea Aublet (1775) and Mariscus Vahl (1805/6) are congeneric. A proposal will be made ep.); Mariscus Vahl is already conserved against 4 ard to add ae to the list of aia rejected in favor of Mariscus (Zarucchi, in pr +Mariscus Scop. (1754) -- Cype he (1982) proposes that Rinorea Aublet be conserved over Conohoria Aublet (see entry no. 36). Index Nominum Genericorum, Farr and cease (1979) list Ropourea Aublet and the replacement names of Scopoli and Schreber as ae of the Verbenaceae rather than the Ebenac [p86I SHNVN OLYANAD S.LATANV THOONUVZ £56 not *Idesia Maxim. (1866) -- Flacourtiaceae. >tRourea is conserved against +Kalawael Adanson (1763). 32 Not Zwingeria Heister ex Fabr. (1759) -- Labiatae; not Zwingera Hofer (1762) -- Nolanaceae. >3Not +Simaruba Boehmer (1760) -- Burseraceae. A schreber (1789), as well as Farr and colleagues (1979), incorrectly attributed the publication of Virecta to Linnaeus filius in the 5 Supplementum Plantarum (1781 [1782]); Virecta was first published by Linnaeus in Plantae Surinamenses (1775). Howa SWith Landia Comm. [ex A. L. Juss.] in synonymy. 5Not +Mappia Heister ex Fabr. (1759) -- Labiatae; not *Mappia Jacq. (1797) -- Icacinaceae. Not *Logania R. Br. (1810) -- Loganiaceae. 8this name of Aublet appears in much of the botanical literature as "Tachigalia." °° Tamonea is accepted as a validly published name of verbenaceous plants. See comments under Fothergilla Aublet (entry no. 56) and in rd (1983) and Stafleu & Cowan (1976). 60h inium Schreber appears as genus number 1545 in the text; in "Addenda et Emendanda" Schreber listed "1545. RHINIUM. Deleatur." Ole ased on genus number citation in the index; not listed in the text. 62Not Bellardia All. (1785) -- Scrophulariaceae. ON ot *Robinsonia DC. (1833) -- Compositae. WOLAYOUUV GCTIONUV AHL JO TVNUNOL VEC C9 “Toa] TABLE 2. Index to generic names listed in Table 1.' Aberemoa Aublet - 1f¢, 25 Abuta Aublet - 3f Acia Schreber - 4, 38 Acioa Aublet - 38 Acouroa Aublet - 5f, 199 Aegiphila Jacq. - 76 Alstonia Scop. - 109 Amasonia L. f. - 171 Ambelania Aublet - 109, 109f Anavinga Adanson - 65 Andira A. L. Juss. - 203 Antirhea Comm. ex A. L. Juss. 75 Apalatoa Aublet - 192 Ardisia Sw. - 63, 63f Aubletia Schreber - 12 Avicennia L. - 59 Bacopa Aublet - 15f Barreria L. - 132f Barreria Scop. - 132 Bellardia All. - 189f Bellardia Schreber - 189 Bellonia L. - 15 Benteca Adanson - 9 Blackstonia Hudson - 93f Blakstonia Scop. - 93 Bockia Scop. - 95 Bonnetia Martius - 73f Bonnetia Schreber - 73 Brami Adanson - 15f Brosimum Sw. - 55, 127 Buchenavia Eichler - 114, Buchnera L. - 129 Caesalpinia L. - 3/7 \ 14F Callicocca Schreber - 28, 53, Calycorectes 0. Berg - 30, Camax Schreber - 52 Carolinea L. f. - 108, 108f Casearia Jacq. - 65, 126, Cassipourea Aublet - 29f Catinga Aublet - 30f Cedrota Schreber - 10 Cephaélis Sw. - 28, 53, 178 Ceranthus Schreber - 87 Chaetocarpus Schreber - 137 Chaetocarpus Thwaites - 137f Chemnicia Scop. - 153 Chesnea Scop. - 28 Chionanthus L. - 8/7, 8/7f Chlora Adanson - 104 Chomelia L. - 75 Clibadium L. - 17 Coccocypselum P. Br. corr. Schreber - 120f, 189 Combretum Loefl. - 23 Comocladia P. Br. - 181 Conohoria Aublet - 36f, 150f Conopea Schreber - 35 Conoria A. L. Juss. - 36 Couepia Aublet - 4, 26f Coumarouna Aublet - 180 Coupoui Aublet - 41f Courimari Aublet - 43f Crudia Schreber - 11, 192 Cryptostomum Schreber - 97 Cuba Scop. - 170 Cucullaria Schreber - 201 Cunninghamia R. Br. - 75f Cunninghamia Schreber - 75 Cyclas Schreber - 11, 192 Cyrtanthus Schreber - 133 Dahuronia Scop. - 92 Dalbergia L. f. - 5, 5f Derris Lour. - 50 Dialium L. - 13 Dickia Scop. - 85 Dimorpha Schreber - 52, 118 Diospyros L. - 115, 152 Dipteryx Schreber - 40, 180 Doliocarpus Rolander - 24, 166 Douglassia Miller - 6f Douglassia Schreber - 6 Dracontium L. - 141 Dugortia Scop. - 117 Duguetia A. St. Hil. - 1, 1f Dupinia Scop. - 176 Duroia L. f. - 41, 41f Ehrhardia Scop. - 6 Eperua Aublet - 118 Ephielis Schreber - 83 Ernstingia Scop. - 83 Evea Aublet - 28, 178 Faramea Aublet - 53 Ferolia Aublet - 117 Fothergilla Aublet - 173f Fothergilla L. - 56f Freyeria Scop. - 87 Galarips Allam. ex L. - 104 Genipa L. - 106, 187 Gentiana L. - 104 SAINVN OFMSNAD S.LATENV ‘THOONUVZ [p861 SEG TABLE 2 (continued). Gerbera L. - 140f Gerberia Scop. - 140 Ghinia Schreber - 173 Ginannia Scop. - 113 Glossoma Schreber - 202 Glossopetalum Schreber - 58 Gnetum L. - 182 Griselinia Scop. - 98 Guatteria Ruiz & Pavon - 1, Gustavia L. - 128, 128f Hambergera Scop. - 23 Hamelia Jacq. - 156 Hedycrea Schreber - 67 Homalium Jacq. - 100, 144 Houmiria A. L. Juss. - 62 Humiria Aublet corr. J. St. Hil. - Hydrolea L. - 157 Hyphydra Schreber - 188 Icacorea Aublet - 63f Idesia Maxim. - 152f Idesia Scop. - 152 Ignatia L. f. - 153 Ilex L. - 72 Ixora L. - 120 Japarandiba Adanson - 128f Jatropha L. - 60 Joncquetia Schreber - 177 Kalawael Adanson - 155f Knoxia L. - 54 Kruegeria Scop. - 204 Labatia Scop. - 72, 137f Labatia Sw. - 72f, 137 Lacis Schreber - 94 Lagunezia Scop. - 144 Landia Comm. ex A. L. Juss. Landolphia P. Beauv. - 109 Lasiostoma Schreber - 153 Legnotis Sw. - 29, 29f Leinkeria Scop. - 154 Lemniscia Schreber - 198 Leonicenia Scop. - 56, 56f Licania Aublet - 26f, 92 Lignonia Scop. - 122 Limborchia Scop. - 48 Linociera Sw. ex Schreber Lita Schreber - 207 Logania R. Br. - 16/7f Loghania Scop. - 167 Lygistum P. Br. - 99 Macanea A. L. Juss. - 70 Macoucoua Aublet - 137f Macrolobium Schreber - 107, Maireria Scop. - 96 Manettia Mutis ex L. - 99 Mapouria Aublet - 112, 162 Mappia Heister ex Fabr. - 166f Mappia Jacq Mappia Schreber - 166 Mariscus Vahl - 148, 148f Mattuschkaea Schreber - 124 Mayepea Aublet - 87f Meisteria Scop. - 110 Melastoma L. - 56, 74, 183, 186 Miconia Ruiz & Pavon - 56, 56f Miegia Schreber - 148 Mnasium Schreber - 146 Moelleria Scop. - 65 Moniera P. Br. - 15f Muellera L. f. - 37 Mussaenda L. - 164 Myristica Gronov. - 200 Myrmecia Schreber - 168 Myrodendrum Schreber - 62 Myrodia Sw. - 140, 140f Myrsine L. - 145 Myrtus L. - 95 Nauclea L. - 106 Nectandra Rolander ex Rottb. Nectris Schreber - 22 Nicandra Adanson - 135f Nicandra Schreber - 135 Oribasia Schreber - 101 Ornitrophe Comm. ex A. L. Juss. - 205 Ouratea Aublet - 105f Outea Aublet - 204 Ozophyllum Schreber - 184 Pacouria Aublet - 9 Palicourea Aublet - 162 Palovea A. L. Juss. - 113 Pamea Aublet - 114f Parinarium A. L. Juss. - 55, 117 Parivoa Aublet - 52 Passoura Aublet - 149 Patabea Aublet - 189 Paullinia L. - 5 Pavetta L. - 76 Payrola A. L. Juss. - 122 Pecheya Scop. - 46 Pekea Aublet - 158 Petaloma Sw. - 95 Petrocarya Schreber - 117 Phyllanthus L. - 33, 89 Picramnia Sw. - 181 Picrium Schreber - 48 Piper L. - 141 Pirigara Aublet - 128f WOLAYOUUV GCTONAUV FHL AO TWNYNOL 0G 69 “10A] Pisonia L. - 59 Plotia Adanson - 145 Ponaea Schreber - 193 Pontopidana Scop. - 44 Porostema Schreber - 103 Portlandia P. Br. - 47 Protium Burm. f Pseudo-Brasilium Adanson - 181 Psychotria L. - 28, 79, 101, 151, 178 Pterocarpus Jacq. - 98, 98f Pterocarpus L. - 98 Quararibea Aublet - 140f Quassia L. - 161 Ramspekia Scop. - 133 Reichelia Schreber - 157 Remirea Aublet - 148f Rhinium Schreber - 185, 185f Rhizobolus Gaertner ex Schreber - 123 Rhopium Schreber - 89 Riana Aublet - 119, 150 Rinorea Aublet - 36, 36f, 119, 149, 150f Rittera Schreber - 134 Robergia Schreber - 155 Robinsonia DC. - 195f Robinsonia Scop. - 195 Rohria Schreber - 179 Rourea Aublet - 155f Ruyschia Jacq. - 167 Salacia L. - 70, 77 Salmasia Schreber - 169 Salmonia Scop. - 201 et - 123 Savastania Scop. - 183 Schwenkfelda Schreber - 156 Schwenkfeldia Schreber - 156 Sciuris Schreber - 147 Sicelium P. Br. - 120, 120f, 189 Simaba Aublet - 14 Simaruba Boehmer - 161f Simira Aublet - 79, 112 Siphonia D. Rich. ex Schreber - 60 Sloanea L. - 2, 12, 43, 43f Smithia Aiton - 139f Smithia Scop. - 139 Solanum L. - 19 Spigelia L. - 91 Saouari Aub Stemodia L. - 85 Stephanium Schreber - 112 Sterbeckia Schreber - 163 Sterculia L. - 66 Strychnos L. - 153 Swartzia Schreber - 134, 194 Swertia L. - 104 Syena Schreber - 86 Symphonia L. f. - 93 Symplocos Jacq. - 31 Talisia Aublet - 143 Tamonea Aublet - 56f, 173f Tapogomea Aublet - 28, 53 Taralea Aublet - 40 Tattia Scop. - 100 Terminalia L. - 175 Ternstroemia Mutis ex L. f. Tetracera L. - 185 Tita Scop. - 29 Tonabea A. L. Juss. - 176 Tonsella Schreber - 190 Tontanea Aublet - 120 Touchiroa Aublet - 11 Trichocarpus Schreber - 2 Trixis Adanson - 17f Trixis P. Br. - 1/7f Trixis Sw. - 17 Uncaria Schreber - 106 Uvaria L. - 25 Vandellia P. Br. ex L. - 85 Vatairea Aublet - 5 Verbena L. - 173 Virecta L. - 164, 164f Vochisia A. L. Juss. - 201 Vochysia Aublet corr. Poiret Vouapa Aublet - 107 Waldschmidtia Scop. - 11 Wernisekia Scop. - 62 Willughbeia Roxb. - 9f Willughbeja Schreber - 9, 9f, 109 Wolffia Horkel ex Schleiden - 131f Wolfia Schreber - 131 Xanthe Schreber - 139 Ximenia L. - 61 Zanonia L. - 42 Zwingera Hofer - 160f Zwingera Schreber - 160 Zwingeria Heister ex Fabr. ‘Except Aublet’s names appearing as main entries and only under that entry number in Table 1. 2Numbers followed by an ‘“f indicate that the generic name is discussed in a footnote to that entry number in Table 1. SANVN OFYANAD S.LATANV THOONUVZ [p861 Eate JOURNAL OF THE ARNOLD ARBORETUM TABLE 3. List of Aublet’s generic names by family.' Anacardiaceae [ANAC] - 177 Annonaceae [ANNO] - +12, +25 Apocynaceae [APOC] - 9, 39, 71, +104, +109 Aquifoliaceae [AQUI] - 72 Asclepiadaceae [ASCL] - 84* Bombacaceae [BOMB] - 108, 140 Bonnetiaceae [BONN] - 26*, 73 Celastraceae [CELA] - 770, ?77, 1747, Chrysobalanaceae [CHRY] - 4, 38, 6/7, 92, 117*, 169 Combretaceae [COMB] - 23, [4+]114, 175 Compositae [COMP] - 17*, 110 Connaraceae [CONN] - *155 Cyperaceae [CYPE] - 78, Dichapetalaceae [DICH] - 179 Dilleniaceae [DILL] - 24, 166, 185* Ebenaceae [EBEN] - 115, 152 Elaeocarpaceae [ELAE] - 2, ?43 Eriocaulaceae [ERIO] - 188 Euphorbiaceae [EUPH] - 8, 33, 34, 60, 69*, 80, 89? Flacourtiaceae [FLAC] - 18, 65, 88, 100, 126, 131, 144 Gentianaceae [GENT] - 48*, 168, 207* Guttiferae [GUTT] - 93, 139*, 196 Haemadoraceae [HAEM] - 208 Humiriaceae [HUMI] - "62", 198 Hydrophyllaceae [HYDR] - 157 Icacinaceae [ICAC] - 132 Iridaceae [IRID] - 32 Lauraceae [LAUR] - 6, 10, 68, 103 Lecythidaceae [LECY] - 42, 44, +128* Leguminosae [LEGU] - +5?, +11, 13, 21, 37, +40, +50, 52, 98, +107, 113, 118*, +134, 2163, 170*, +180, +192, +194, 199, +203, +204* Loganiaceae [LOGA] - 91, 135, 153 Lythraceae [LYTH] - 49 Marcgraviaceae [MARC] - 102, Mayacaceae [MAYA] - 86 Melastomataceae [MELA] - #56, 74, 95, 183, 186, 191, 202 Meliaceae [MELI] - 27 Menispermaceae [MENI] - 3* Monimiaceae [MONI] - 165 Moraceae [MORA] - 16, +55, 81, 125, +127 Myristicaceae [MYRI] - 200 Myrsinaceae [MYRS] - +63, 145 Myrtaceae [MYRT] - 30*? Nyctaginaceae [NYCT] - 59 Ochnaceae [OCHN] - *105 Olacaceae [OLAC] - 61, 90 Oleaceae [OLEA] - +87 Passifloraceae [PASS] - ?159 Piperaceae [PIPE] - 141 Podostemonaceae [PODO] - 94 Polygalaceae [POLY] - 97 Proteaceae [PROT] - 154 Quiinaceae [QUIT] - 142, 195 Rapateaceae [RAPA] - 146 Rhizophoraceae [RHIZ] - 29 Rubiaceae [RUBI] - 7, 20, +28, (+)41, 46, 47, +53, 54*, 75, 79, 99%, 101*, +106, 111, 112, 120, 121, 124, 133, 151*, 156*, 162, 164, = 67 +178*, 187, 189 Rutaceae [RUTA] - 57, 147, 184 Sapindaceae [SAPI] - 51, 83, 143, 172, 193, 205 Sapotaceae [SAPO] - 137 Scrophulariaceae [SCRO] - *15, 35, 85, 129 Simaroubaceae [SIMA] - 14, 160, *161, 181 Solanaceae [SOLA] - 19 Sterculiaceae [STER] - 66 Symplocaceae [SYMP] - 31 Theaceae [THEA] - +176* Tiliaceae [TILI] - 12* Trigoniaceae [TRIG] - 197* Turneraceae [TURN] - 130 Verbenaceae [VERB] - 76*, +171, 173 [VOL. 65 1984] ZARUCCHI, AUBLET’S GENERIC NAMES 239 TABLE 3 (continued). Violaceae [VIOL] - (+)36, 119, 122, Family uncertain - ?43, 270, ?77, 149, (*)150 2159 Vochysiaceae [VOCH] - 138*, "201" "y listing by family of Aublet's 208 generic names treated in Table 1. The numbers indicate their current taxonomic acceptance (underlined) or treatment as a4 synonym (without underlining). Numbers corresponding to polytypic genera are followed by an asterisk (*), those of questionable status are followed by a question mark (?), and those accepted with a modified orthography appear in quotes ("). The five genera that are questionably placed in a family are both underlined and preceded by a question mark (?). Symbols preceding a generic number indicate a conserved name (*), a rejected name (+), a later homonym (#), or a name that was superfluous when published (+). Parentheses or brackets around a symbol indicate that a proposal to conserve/reject has been (*) or should be [*] made. (excluding main-entry names of Aublet) given in TABLE | and its footnotes. inally, a review of the currently accepted taxonomic status for Aublet’s genera, grouped by families, is presented in TABLE 3. The orthography of taxonomic names varied frequently within and among the individual works of Aublet’s time. Spelling variants and their location are listed in TABLE |. Scopoli (1777) used ‘“*V” for both modern-day ““U” and “V,” and these variants are presented in quotation marks in TABLE |. Article 73.5 of the Sydney Code (Voss et a/., 1983) has recently established that the use of “Vv” in this situation should not be considered nomenclaturally distinctive. In Aublet’s Histoire des Plantes de la Guiane Francoise, descriptions of representative species of his new genera were accompanied by separate line rawings. Most of these new genera (177 of 208) were monotypic, although others contained up to 6 species. The 31 polytypic genera described by Aublet (with the number of included species) are: Abuta (2), Apeiba (4), Baillieria (2), Caraipa (4), Catinga (2), Coussapoa (2), Coutoubea (2), Faramea (2), Goupia (2), Icica (6), Mabea (2), Manabea (3), Matelea (2), Nacibea (2), Nonatelia (6), Parinari (2), Parivoa (2), Pekea (2), Pirigara (2), Qualea (2), Quapoya (2), Ronabea (2), Sabicea (2), Saouari (2), Tachigali (2), Taonabo (2), Tapogomea (5), Tigarea (2), Trigonia (2), Vouapa (2), and Voyria (2). In his Introductio ad Historiam Naturalem, Scopoli (1777) considered only 149 of Aublet’s generic names and proposed replacements for 44. He accepted 90 genera and placed 13 in synonymy under previously described genera; Z genera were treated as synonyms of other names of Aublet. Several of these replacement names were retained by Schreber (1789, 1791) in his edition of the Genera Plantarum. Others, however, were disregarded in favor of Schre- ber’s own new names. In several cases Schreber took up names proposed by Swartz in his Prodromus (1788) or by Linnaeus fi/ius in his Supplementum 240 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Plantarum (1781). In addition, Schreber totally ignored 88 of Aublet’s generic names. Together, Scopoli and Schreber considered 110 of Aublet’s genera and ignored 49, The Genera Plantarum by A. L. de Jussieu (1789) accounted for all of Aublet’s 208 Neotropical genera of plants. For 8 genera, Jussieu proposed replacement names that were quite similar to Aublet’s names; an additional 8 names were simply orthographic variants. Jussieu accepted a total of 182 of Aublet’s genera, including the 8 similar replacement names. Twenty-six genera were considered to be taxonomic synonyms, with 8 of these synonyms of other genera of Aublet. Jussieu placed all except 20 genera in his “natural system’ of classification. The genera described by Aublet (1775) fall into 79 modern plant families if one views the questionably placed Senapea as a genus of Passifloraceae and if one accepts the Cecropiaceae and Goupiaceae. The best-represented families are the Rubiaceae and Leguminosae, with 26 and 21 of Aublet’s genera at- tributed to them, respectively. No fewer than 9 accepted higher-plant family names are based on Aublet’s generic names: Cabombaceae, Goupiaceae, Hu- miriaceae, Mayacaceae, Quiinaceae, Rapateaceae, Simaroubaceae, Trigoni- aceae, and Vochysiaceae. Many of Aublet’s generic names have been the subject of conserva- tion/ rejection proposals. S1x of Aublet’s names have been conserved (see TABLE 1, items 15, 62, 105, 155, 161, and 201). Two of these, Houmiuri (62) and Vochy (201), are conserved with altered orthographies— Humiria and Vochysia. Hekking (1982) proposes to conserve one of Aublet’s violaceous generic names, Rinorea (150), over another, Conohoria (36). Ouratea (105) has been con- served, but no name is listed as being rejected in its favor (Voss et al., 1983). Twenty-six of Aublet’s generic names have been rejected, most of them apparently in favor of a more widely used or preferred name. Two of the rejected genera, Pacouria (109) and Vouacapoua (203), are considered to be taxonomically distinct from their conserved counterparts; two additional gen- era, Aberemoa (1) and Acouroa (5), may also fall into this category. During the course of the present investigation, it has come to my attention that three more of Aublet’s genera need to be considered for rejection based on the identifi- cations cited by Howard (1983). Coupoui (41) will be proposed for rejection in favor of Duroia L. f., and Remirea (148) in favor of Mariscus Vahl (Zarucchi, in prep.). Pamea (114), a monotypic genus when described by Aublet, 1s listed by Howard as a species of Buchenavia Eichler (1866) (Combretaceae), based upon a personal communication from Dr. C. Stace; 1t appears that a conser- vation/rejyection proposal is in order. Another problem surrounds the correct identification of Catinga moschata Aublet, the generic econ ps a coe (McVaugh, 1956), which has been questionably identified as Ca Sandw. If Cal/ycorectes O. Berg (1856) and Catinga prove to be congeneric, a rejecuon proposal may be desirable. Two of Aublet’s generic names, Ore/ia (104) and Pirigara (128), were ille- gitimate when published because they included the types of previously de- scribed genera. The description of Orel/ia Aublet contained the earlier-published 1984] ZARUCCHI, AUBLET’S GENERIC NAMES 241 Allamanda L. (1771), and the protologue of Pirigara included Japarandiba, a Marcgrave name validly published by Adanson in 1763. Japarandiba is rejected in favor of Gustavia L. (1775). Fothergilla (56) is the only name used by Aublet that is a later homonym (of the hamamelidaceous Fothergilla L. (1774)) and therefore not acceptable. Of the remaining 167 genera, 52 are considered to be taxonomic synonyms of previously published genera and 115 are accepted. Only 7 of these 115 genera are accepted with reservation as to placement to family (43, 70, 77, 159, 163) or as to recognition as generically distinct within a family (89, 174). Although current taxonomic opinion varies on the acceptance or rejection of some of Aublet’s generic names, a large number of them are valid and in use today. The many unusual-sounding names published more than two cen- turies ago continue to fascinate, if not tongue-tie, new generations of tropical American botanists. ACKNOWLEDGMENTS I wish to express my gratitude to the staffs of the Department of Botany, British Museum (Natural History), London, and the Royal Botanic Gardens, Kew, England, where this research was initiated under a NATO Postdoctoral Fellowship in Science. Valuable bibliographic assistance was provided by the librarians of these institutions and of the Linnean Society of London. I also wish to thank the staff of the Department of Botany of the Smithsonian Insti- tution, particularly D. H. Nicolson. Additional work was carried out under a Smithsonian Fellowship. LITERATURE CITED Aus.et, F. 1775. Histoire des plantes de la Guiane francoise. 4 vols. Didot, Paris. Barnesy, R. C., & B. A. Kaukorr. 1971. Supplementary notes on American Meni- spermaceac. VIII. A generic survey of the American Triclisicae and Anomosper- meae. Mem. New York Bot. Gard. 22(2): 1-90. Farr, E. R., J. A. Leussink, & F. A. STAFLEU. 1979. a nominum genericorum (plantarum). 3 vols. Bohn, Scheltema & Holkema, Utrec HEKKING, W.H. A. 1982. Proposal to conserve 5262 ae Aublet be against multaneously published Conohoria Aublet (1775) (Violaceae). Taxon 31: , 755. bance R.A. 1983. The plates of Aublet’s Histoire des plantes de la Guiane rte J. Arnold Arbor. 64: 255-292. Jussieu, A. L. pe. 1789. Genera plantarum. Herissant, Pari Kusitzki, K. 1978. Caraipa and Mahurea (Bonnetiaceae). ie B. Macuir_, ed., The otany of the Guayana Highland, part X. Mem. New York Bot. Gard. 29: 82-138. LINNAEUS, C. von. 1781 [1782]. Supplementum plantarum. Braunschweig. LuNDELL, C. L. 1981. Neotropical Myrsinaceae— VI. Phytologia 49: 341-354 McVauau, R. 1956. Nomenclatural notes on Myrtaceae and related families. Taxon 5: 133-147. Necker, N. J. pe. 1790. Elementa botanica. 3 vols. Neuwied. 242 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Scureser, J. C. D. von. 1789, 1791. Genera plantarum. ed. 8. 2 vols. Varrentrapp & Scopou, G. A. 1777. Introductio ad historiam naturalem. eeu a Prague. STAFLEU, F. A. 1971. Linnaeus and the Linnaeans. A. Oosthoek, Ut R. S. Cowan. 1976, 1981. Taxonomic literature. ed. 2. eH i. 3. Bohn, Scheltema & Holkema, Utrecht SteARN, W. T. 1976. Union of Chionanthus and Linociera (Oleaceae). Ann. Missouri Swartz, O. P. 1788. Nova genera & species plantarum seu prodromus. Stockholm. Voss. E. G., et al. 1983. International code of botanical nomenclature [Sydney Code]. Bohn, Scheltema & Holkema, Utrecht. DEPARTMENT OF BOTANY Current address SMITHSONIAN INSTITUTION MISSOURI haere GARDEN WASHINGTON, D. C. 20560 P. O. Box St. Louris, MissourR! 63166 1984] BAAS, ILEX COLLINA AND NEMOPANTHUS 243 VEGETATIVE ANATOMY AND THE TAXONOMIC STATUS OF ILEX COLLINA AND NEMOPANTHUS (AQUIFOLIACEAE) PIETER BAAS In 1974 Clark clarified part of the confusion in the literature with respect to three deciduous North American species of //ex L.: I. longipes Chapman, /. decidua Walter, and J. collina Alexander. Because /. collina has apopetalous corollas, calyx lobes that are semipersistent on the fruits and inconspicuous in staminate flowers, and free stamens, he transferred it to Nemopanthus Raf., noting that J. /ongipes as understood by most authors should be treated as a synonym of Nemopanthus collinus. In my accounts of the vegetative anatomy of the Aquifoliaceae (Baas, 1973, 1975, 1978), I noted the close anatomical similarities of Nemopanthus with several species of //ex from subg. PRINUS (L.) Maxim. and concluded (1975, p. 355) that merging Nemopanthus with Ilex “would not meet with any opposition from vegetative anatomists,” the peculiar cuticular markings of Nemopanthus leaves being the only distinguish- ing anatomical character. Dudley and Eisenbeiss (U. S. National Arboretum) drew my attention to Ilex collina and expressed their view that the transfer to Nemopanthus by Clark might after all have been unjustified. In view of the desirability of a proper understanding of the affinities and taxonomic status of this species as well as of Nemopanthus mucronatus (L.) Trel., and also because of the potential or actual importance of these species in horticulture, leaf and xylem anatomy of Ilex collina was studied in considerable detail, and additional specimens of Nemopanthus mucronatus were examined in order to check the constancy (and hence the diagnostic and taxonomic value) of its anatomical characters. MATERIALS AND METHODS The techniques employed have been described earlier (Baas, 1973, 1975). Sections and macerations were studied by light microscopy, and gold-coated surfaces of critical-point dried, previously FAA-fixed material were examined with a scanning electron microscope. All material used for this study was from the herbarium or the living collections of the U. S. National Arboretum, Wash- ington, D. C. The comparative part of the work is largely based on material from the Rijksherbarium collections, detailed previously (Baas, op. cit.). © President and Fellows of Harvard College, 1984. Journal of the Arnold Arboretum 65: 243-250. pee 1984. 244 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 RESULTS ILEX COLLINA LEAF ANATOMY Material studied. U.S.A. Virginia: Mountain Lake, Shanks s.n. (NA). West Virginia: Cheat R., Sharp 1070 (NA); Cheat Mt., Hutton s.n., 13 mee: oo (NA). Pennsylvania: cult. rbor. Barnes Foundation, aes Fogg s.n. (NA). Washington, D. C.: cult. from seed collected in 1975 at type locality, U. S. National i sretAs 38173 (NA); from seed collected in 1974, Dudley & Tisenbeiss 400 1978 (NA); from seed collected in 1975, U.S. National . Aberin s.n. (alcohol-preserved material; 1 Surface view. Long, unicellular, partially septate hairs usually present, especially on veins and abaxial surface (FIGURE 1). Cuticle of adaxial and abaxial surfaces smooth to rather faintly striate (FIGURES 4, 5). Epidermal cells of adaxial surface with straight to (especially in leaves of juvenile specimens) undulating anticlinal walls, those of abaxial surface with strongly undulating (zigzag pattern) walls. Epidermal cell pattern modified over major veins and midrib only. Stomata confined to abaxial surface, irregularly anomocytic to cyclocytic with (3 or) 4 to 7 (to 10) neighboring/subsidiary cells (FiGuRE 2), occasionally resembling anisocytic or laterocytic stomata (cf. Den Hartog-Van Ter Tholen & Baas, 1978), guard cell pairs (21-)26-32(-35) by (18-)20-26(-29) «wm. Peristomal rims and cuticular T-pieces absent. Transverse section. Lamina dorsiventral, 80-140 wm thick. Cuticle ca. 1 wm thick. Unspecialized epidermal cells usually flattened, the adaxial cells much larger than abaxial, often bulging, rarely with periclinal wall divisions and/or mucilaginous. Hypodermis absent. Mesophyll composed of | (rarely 2) layers of short to tall palisade cells and fairly loose spongy tissue. Midrib grooved adaxially, prominently raised abaxially, supplied with single shallowly arc- shaped collateral vascular bundle; vascular bundle with abaxial cap of thin- walled sclerenchyma fibers, very rarely (only in Hutton s.n.) with incurved margins. Ground tissue of midrib parenchymatous to collenchymatous. Veins mostly embedded, with parenchymatous bundle sheath poorly differentiated and extending to upper and lower epidermis only in major veins. Leaf margin with chlorenchymatous ground tissue. Petiole vascularization similar to that of midrib, proximal part with single bundle (type |: cf. Baas, 1975), distal portion with additional small latero-dorsal wing bundles (types 3 and 4). Crys- tals present as druses especially in vicinity of vascular bundles. Woop ANATOMY This description is based on a stem 6 mm in diameter from an alcohol- preserved specimen cultivated in the U.S. National Arboretum. Quantitative values should be considered with caution because the specimen was from an immature plant (3 years old). The presence of spiral thickenings was checked 1984] BAAS, ILEX COLLINA AND NEMOPANTHUS 245 10084, jon Ficures 1-3. 1, 2, Mex collina: 1, unicellular, septate hair; 2, anomocytic (left) and irregularly cyclocytic (right) stomatal complex. 3, Nemopanthus mucronatus, cyclocytic, anomocytic, and laterocytic stomata. and confirmed in thin twigs of all herbarium specimens used in studying the leaf anatomy (see above). Wood semi-ring porous. Growth rings distinct. Vessels ca. 160 per mm’, solitary (ca. 30%) and in radial multiples of | to 6, angular to oval in transverse section, tangential diameter (17-)27(-40) um, vessel member length (270-)540(-690) um. Intervessel pits mainly opposite (some transitional or tending to alternate), oval or elongate, 5-8 um in horizontal diameter, with slitlike apertures enclosed within pit borders. Vessel—ray and vessel—paren- chyma pits similar but half bordered. Perforations scalariform in oblique end walls with (11-)14(-20) bars. Spiral thickenings well developed (FiGuRE 6). Vessel contents and tyloses absent. Ground tissue composed of thin-walled fiber-tracheids (570-)7 10(-870) nm long with bordered pits most numerous on radial walls, the pits 5-6 um in diameter, with slitlike apertures extending only slightly beyond pit borders. Spiral and annular thickenings prominent (FIGURE 7). Parenchyma diffuse, diffuse in small aggregates, and rarely scanty paratra- cheal, in strands of (3 or) 4 to 6 (or 7) cells. Rays heterogeneous II, uniseriate rays ca. 8 per mm, multiseriate rays 3 (or 4) cells wide and ca. 4 per mm, tallest rays up to 0.6 mm high. Sheath cells and crystals absent. Judging from comparisons of juvenile and mature secondary xylem in other Ilex species, one would expect mature wood of Ilex collina to have considerably broader rays, somewhat fewer vessels, and somewhat wider and longer axial elements. Qualitative or even major quantitative differences with the above description are unlikely, however. Ficures 4-10. 4-7, Ilex collina: 4, smooth abaxial cuticle; 5, finely striate cuticle: , secondary xylem vessel with helical thickenings; 7, fiber-tracheid with spiral to helical thickenings. 8-10, Nemopanthus mucronatus: 8, strongly ridged cuticle with fine wax 1984] BAAS, ILEX COLLINA AND NEMOPANTHUS 247 NEMOPANTHUS MUCRONATUS VEGETATIVE ANATOMY! Material studied.2 CANADA. Nova Scotia: Gorham 45.1351 (L). U.S.A. Maine: Hancock Co., Dudley 8031 (L). New Hampshire: Schweinfurth & Correll 11235 (L). Pennsylvania: Pike Co., Brumbach 7982 (1); cult. Arbor. Barnes Foundation, Merion, Fogg 5.n. (1). Surface view. Glabrous (except in Schweinfurth & Correll 11235, which has few short, unicellular hairs on adaxial side of midrib). Cuticle of adaxial surface smooth or faintly to conspicuously striate, that of abaxial surface with con- spicuous, rather widely spaced ridges (FiGureE 8). Epidermal cells of adaxial surface with straight to curved (to undulating) walls, those of abaxial epidermis with curved to strongly undulating walls. Epidermal cell pattern modified over major and most minor veins (cells elongate parallel to venation). Stomata confined to abaxial epidermis, irregularly cyclocytic (FIGURE 3), sometimes tending to anomocytic or laterocytic with (3 or) 4 to 6 (or 7) subsidiary cells, guard cell pairs (18—)2 1—25(-30) by (1 7-)18-21(-26) um. Peristomal rims pres- ent. Cuticular T-pieces absent. Transverse section. Lamina dorsiventral, 90-190 um thick. Cuticle usually ca. 1 (rarely up to 2) um thick. Unspecialized epidermal cells usually flattened, the adaxial cells larger than abaxial, often with convex inner periclinal walls and periclinal wall divisions, only rarely resembling cells. Hypodermis absent. Mesophyll composed of | layer of palisade cells and pat loose spongy tissue. Midrib very shallowly grooved to raised adaxially, prominently raised abaxially, supplied with single shallowly arc-shaped collateral bundle (some- times with additional small dorsal bundle) having poorly differentiated abaxial sclerenchyma cap. Ground tissue of midrib collenchymatous. Veins mostly vertically transcurrent through parenchymatous to collenchymatous bundle- sheath extensions, only smallest veins embedded in mesophyll. Petiole with single collateral bundle having incurved margins at basal end (type 2; cf. Baas, 1975) and additional latero-dorsal wing bundles at distal end (type 4). Crystals present as druses, especially in vicinity of vascular bundles. ma | WooD ANATOMY The wood specimens described in Baas (1975), as well as twigs of the her- barium specimens cited above, were examined for presence or absence of spiral thickenings. Such tertiary thickenings were absent from the fiber-tracheids in 'An amplified, nearly identical version of the previously published account (Baas, 1975). 2In addition to that studied previously, listed in Baas (1973, 1975). platelets; 9, secondary xylem vessel without spiral thickenings; 10, detail of vessel tip with very faint wall thickenings. 248 JOURNAL OF THE ARNOLD ARBORETUM TABLE |. [VOL. 65 Differences between Ilex collina and Nemopanthus mucronatus. CHARACTER SPECIES Ilex collina Nemopanthus mucronatus Leaf Abaxial cuticle Indumentum Stomata Average length of guard cell pairs (um Midrib Veins ae ae Smooth or striate Long, partly septate hairs Strongest tendency toward anomocytic 6-32 Typically ae es with eveloped collen- prominent in epidermal cell pattern Prominent in fiber-tracheids Characteristically ridged Usually a ; if present, airs short, nonseptate Strongest tendency toward cyclocytic 21-25 Usually raised, with well- developed collenchyma in round tissue Mostly vertically transcur- rent, prominent in epi- dermal cell pattern as et absent, or very in secondar and vessels t and in vessels only aie all material studied (see Ficure 9), although Dudley 8031 and Schweinfurth & Correll 11235 showed extremely fine thickenings in some vessel members, especially near the tails (FiGuURE 10). No other material studied had any spiral thickening in the vessels (see FiGure 9). COMPARISONS AND DISCUSSION ILEX COLLINA AND NEMOPANTHUS MUCRONATUS From the above descriptions many similarities in general characters are apparent between the two species. The differences worthy of listing are shown in TABLE 1. Most of them (indumentum, stomatal type and size, midrib and vein characters, spiral thickenings in the wood) are only of interest as diagnostic aids at the species level. However, when these differences are considered to- gether with the more significant one of cuticular sculpturing, they constitute enough evidence to render questionable Clark’s (1974) transfer of Ilex collina to Nemopanthus. Before any decision can be made, it must first be determined whether //ex collina has closer anatomical affinities with other //ex species than with Nemopanthus mucronatus. ILEX COLLINA AND OTHER ILEX SPECIES The leafand xylem anatomy of //ex collina, as well as its macrom orphological characters and its deciduous habit, are quite similar to those of several species grouped together in subg. Prinus by Loesener (1942). In fact, in leaf and wood 1984] BAAS, ILEX COLLINA AND NEMOPANTHUS 249 anatomy a species like /. decidua is much more similar than Nemopanthus mucronatus to I. collina; the only differences are in some minute details of the indumentum and in the outline of the abaxial epidermal cells. I also studied two specimens labeled J. /ongipes Chapman in the Rijksherbarium (U.S.A.: Massachusetts, cult. Arnold Arboretum 22659 (Boom 39981); Tennessee, Bilt- more Herb. 4063). Apparently these specimens are not /. co//ina because their indumentum is different (short, unicellular hairs instead of long ones; see also description in Baas, 1975). They also differ from the J. decidua specimen that I studied (1975); this might have been expected considering Clark’s remark that J. /Jongipes sensu Trelease in fact belongs to the 7. decidua complex. Ap- parently identifications on herbarium labels in this group of //ex species are far from reliable, and I therefore refrain from earmarking a particular //ex species as being anatomically closest to /. co/lina. For the present problem, it is significant only that several species of subg. PRiNuS “ser. B’? PRINOIDES anatomically resemble /. co/lina more closely than J. collina resembles Nemo- panthus mucronatus. Ilex serrata Thunb. and /. verticillata (L.) A. Gray of ‘*ser. A’? EUPRINUS (sensu Loesener) (= PRINus) also resemble J. collina in leaf anatomy but differ markedly in their wood anatomy because they lack spiral thickenings in the fibers and vessels. NEMOPANTHUS MUCRONATUS AND ILEX As stated before, the only constant anatomical difference between //ex and Nemopanthus mucronatus is the sculpturing of the abaxial cuticle. Within //ex (especially in some tropical species) sculpturing ranges from smooth to con- spicuously striate, but it is nowhere near as marked as in Nemopanthus. Another distinguishing character of N. mucronatus 1s its lack of conspicuous spiral thickenings, which are typical for all //ex species studied from temperate regions except J. serrata and J. verticillata. It is clear that although cuticular sculpturing is of considerable diagnostic value, it alone cannot justify the separation of two obviously closely related genera. All characters must be examined to determine whether other differences are sufficient to maintain Nemopanthus as a separate genus. SUGGESTIONS FOR TAXONOMIC TREATMENT Ilex collina is anatomically much closer to several //ex species belonging to “ser. B” PrinorpDes of subg. Prinus than it is to Nemopanthus mucronatus. Translated into taxonomic practice, this would favor the reinstatement of Nemopanthus collinus in Ilex, in contrast to Clark’s (1974) treatment of the species. Nemopanthus mucronatus shows anatomical affinities to several //ex species of ‘“‘ser. A’? EUpRINus rather than to those of “‘ser. B’’ PRINOIDEs sensu Loesener. If it is true that the floral characters of J. co/lina break down the boundaries between //ex and Nemopanthus, this implies that N. mucronatus should be transferred to //ex and should in the future be treated as J/ex mucronata. However, if floral and other macromorphological distinctions remain, the ad- ditional differences in cuticular structure can be used 1n support of maintaining Nemopanthus as a monotypic genus. 250 JOURNAL OF THE ARNOLD ARBORETUM VOL. 63 ACKNOWLEDGMENTS I am greatly indebted to Theodore agai) and Gene Eisenbeiss (U. S. Na- tional Arboretum, Washington, D. C.) for the generous supply of research material and for drawing my attention to this interesting problem. I also thank Paul van Veen for preparing numerous anatomical slides of troublesome her- barium material. LITERATURE CITED Baas, P. 1973. The wood anatomical range in aa and its ecological and phylogenetic significance. Blumea 21: 193-2 75. Vegetative anatomy and the affinities oe Aquifoliaceae, Sphenostemon, Phelline and ar ee Ibid. 22: 311-407. : 8. Inheritance of foliar and nodal anatomical characters in some J//ex hy- he J. Linn. Soc., Bot. 77: 41-52 CLark, R. C. 1974. Ilex collina. A second species of Nemopanthus in the southern Appalachians. J. Arnold we 435-441. HarTOG-VAN TER THOLEN, R. M. DEN, & P. Baas. 1978. Epidermal characters of the Celastraceae sensu lato. Acta Bot. Neerl. 27: 355-388. LOESENER, T. 1942. Aquifoliaceae. Jn: A. ENGLER & K. PRANTL, eds., Nat. Pflanzenfam. ed. 2. 20b: 36-86. RIJKSHERBARIUM Box 9514 2300 RA LEIDEN, THE NETHERLANDS 1984] SCHLARBAUM ET AL., METASEQUOIA AND SEQUOIA — 251 THE CHROMOSOMES AND RELATIONSHIPS OF METASEQUOIA AND SEQUOIA (TAXODIACEAE): N UPDATE! S. E. SCHLARBAUM, T. TSUCHIYA, AND L. C. JOHNSON In 1948 Stebbins reported on the chromosome number and relationships of the newly discovered Metasequoia glyptostroboides Hu & Cheng (2n = 2x = 22) and Sequoia sempervirens (D. Don) Endl. (2n = 6x = 66). Further research on the karyological relationships between these two taxodiaceous species has not been reported in the following years. The observation of marker chro- mosomes with unusual structures in various species of Taxodiaceae (Schlar- baum & Tsuchiya, 1975, 1976, 1981) stimulated the authors to investigate the chromosome morphology of Metasequoia and Sequoia for possible indications of cytotaxonomic and evolutionary relationships. This paper is a brief report addressing only the marker chromosome differences between Metasequoia and Sequoia. Expanded accounts of the overall karyotypic features of these species will be published elsewhere (Schlarbaum & Tsuchiya, 1984; Schlarbaum et ai/., 1983). Interest in the chromosome complement of Sequoia developed after Lawson (1904) suspected that the species was polyploid. The chromosome number of Sequoia was determined with certainty by the independent meiotic studies of Hirayoshi and Nakamura (1943) and Stebbins (1948). FiGure | is the previ- ously unpublished photo by Hirayoshi and Nakamura (both now deceased) showing the chromosome configurations from which the hexaploid count was determined. As shown by this photograph and the drawings of microsporocytes by Stebbins (1948) and Hirayoshi and Nakamura (1943), multivalents are present, suggesting that Sequoia is at least partially autopolyploid. Before the discovery of Metasequoia was published, some scientists (Jen- sen & Levan, 1941; Hirayoshi & Nakamura, 1943) speculated whether Se- quoiadendron giganteum (Lindley) Buchholz, previously classified as Sequoia gigantea (Lindley) Decne., may have contributed one or more genomes to Sequoia. The discovery of living Metasequoia trees naturally stimulated hy- potheses involving ancestral relationships between Metasequoia and Sequoia. These inferences were based upon the morphological similarities between the two species, and probably in part on the previous confusion of fossil remains of Sequoia and Metasequoia (Miki, 1941; Chaney, 1951). After determining the diploid nature of Metasequoia, Stebbins (1948) considered the plausibility ‘Contribution from the oe DEAE OnOmy, poled State University, Publ ished with the approval of the Director of the I 2769. © President and Fellows of Harvard College, 1984. Journal of the Arnold Arboretum 65: 251-254. April, 1984. 252 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 ed £9 s 4 Go » "Bae Ficure |. Microsporocyte of Sequoia sempervirens showing complete chromosome complement (2n = 2.x = 66) and various multivalents, x ca. 2200 (Hirayoshi & Naka- mura, unpubl.). ofa Metasequoia species being a Mesozoic or early Tertiary ancestor of Sequoia. More recently, Saylor and Simons (1970) speculated on the possibility of chro- mosomal relationships between Mefasequoia and Sequoia. In the present study, observations for marker chromosomes were made on the root-tip mitosis of Sequoia and Metasequoia. Sequoia seeds from an un- known locality were obtained from a commercial company and germinated. Metasequoia cuttings were taken from trees originating from the only extant population in western Hubei Province, China, and rooted. Cytological meth- odology followed Schlarbaum and Tsuchiya (1976). Vouchers are deposited at CS. : Our karyological observations show the presence of marker chromosomes with unusual structures in Metasequoia and Sequoia (FiGuRE 2, a, b). Meta- sequoia has three pairs of chromosomes with long kinetochore regions (FIGURE 2, a). Sequoia has two pairs of SAT-chromosomes with long secondary con- strictions in which chromomeres are visible, and another chromosome pair that probably has microsatellites attached to the short arm (Schlarbaum, 1980; Schlarbaum & Tsuchiya, 1984) (Figure 2, b; microsatellites not visible). 1984] SCHLARBAUM ET AL., METASEQUOIA AND SEQUOIA — 253 nw lal mA f | mm WAC Ficure 2. Marker chromosome pairs, x 3000: a, Metasequoia Epona: 2n= 2x = 22, showing long kinetochore regions; b, Sequoia sempervirens, 2n = 6x = 66 (left and center pairs show long secondary constrictions and corresponding satellite’ bodies). CONCLUSIONS The marker chromosome type found in Metasequoia was not observed in Sequoia, indicating the lack of a direct cytotaxonomic relationship between the two species, although the effects of amphiplasty (Navashin, 1928) cannot be entirely discounted. Based on the cytological evidence presented, genomic contribution of the extant Metasequoia species to the polyploidy of Sequoia 1s considered unlikely. It must be recognized that the chromosome studies were conducted on monotypic, relictual genera that undoubtedly contained more species in past geologic epochs. However, the fossil record supports the cyto- logical conclusion that Sequoia was differentiated independently from Meta- sequoia but probably came from the same general ancestral stock (Arnold & Lowther, 1955). LITERATURE CITED ARNOLD, C. A., & J. S. LowtHer. 1955. A new Cretaceous conifer from northern Alaska. Amer. J. Bot. 42: 522-528. 254 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 CHANEY, R. W. 1951. A revision of fossil Sequoia and Taxodium in western North America based on the recent discovery of Metasequoia. Trans. Amer. Philos. Soc., n.s. 40: 169-263. a I., & Y. NAKAMURA. 1943. Chromosome number of Sequoia sempervirens. t. & Zool. 2: 73-75. ae H., & A. Levan. 1941. Colchicine-induced tetraploidy in Sequoia gigantea. Hereditas 27: 220-224. Lawson, A. A. 1904. The gametophyte, ete abo fertilization, and embryo of Se- quoia sempervirens. Ann. Bot. (London) 18: 1-28. Miki, S. 1941. On the change of flora in ee Asia since Tertiary period (I). The clay or lignite beds’ flora of Japan, with special reference to the Pinus trifolia beds in Central Hondo. Jap. J. Bot. 11: 237-303. NavasHIN, M. 1928. ‘“‘Amphiplastie’’—eine neue karyologische Erscheinung. Proc. Int. Conf. Genet. 5: 1148-1152. SAYLOR, - C., & H. A. Simons. 1970. Karyology of Sequoia sempervirens: karyotype eee chromosomes. Cytologia 35: 294-303. SCHLARBAUM, S. E. 1980. Cytotaxonomic relationships within Taxodiaceae. 217 pp. npubl. D. dissertation, Colorado State University, Fort Collins, Colorado. my bras NSON, & T. Tsucuiya. 1983. Chromosome studies of Metasequoia ebprosrobordesand Taxodium distichum. Bot. Gaz. (Crawfordsville) 144: 559-565. HIyA. 1975. The chromosome study of giant sequoia, Sequoiadendron aes ‘Silvae Genet. 24: 23-26. & ——. | 76. Chromosome study of Japanese umbrella pine. J. Hered. 67: 65-67. & ———. 1981. Differential reactivity in tree chromosome staining. /bid. 72: 62, 63. A ee = of coast redwood, Sequoia sempervirens on) Endl. Silvae Genet. (in Rese G. L., Jr. 1948. The ae nee and relationships of Metasequoia and Sequoia. Science 108: 95-98. LES. T. T. DEPARTMENT OF uel DEPARTMENT OF AGRONOMY UNIVERSITY OF TEN COLORADO STATE UNIVERSITY KNOXVILLE, ane SSEE 47901. 1071 Fort Coiuins, COLORADO 80523 L.C. J. STATE AND PRIVATE FORESTRY, 1992 FoLweLL AVENUE St. PAUL, MINNESOTA 55108 iS ICSEB — Ill THIRD INTERNATIONAL CONGRESS OF SYSTEMATIC AND EVOLUTIONARY BIOLOGY, 1985 The Congress will be held on 4-10 July 1985 at the University of Sus- sex, near Brighton, England. The following Congress Symposia are being organized: Symbiosis in Evolution Conservation of Tropical Ecosystems Biogeographic Evolution of the Malay Archipelago Adaptational Aspects of Physiological Processes Co-evolution in Ecosystems and the Red Queen Hypothesis Angiosperm Origins and the Biological Consequences The Measurement of Rates of Evolution Molecular Biology and Evolutionary Theory Co-evolution and Systematics Molecules vs. Morphology in Phylogeny: Conflict or Compromise? Random and Directed Events in Evolution Biochemical Innovation in Microbial Communities There will also be Special Interest Symposia on other topics, as well as sessions for contributed papers, films, and poster papers. For further information write to: Professor Barry Cox ICSEB Congress Office 130 Queen’s Road Brighton, Sussex BN1 3WE England Journal of the Arnold Arboretum April, 1984 CONTENTS OF VOLUME 65, NUMBER 2 Fruits and Seeds of the Cunoniaceae. WILLIAM C. DICKISON 4.63605 s54 004005 ev eaes banc veadevibeness 149 Anatomy of the Palm Rhapis excelsa, X. Differentiation of Stem Conducting Tissue. P. B. TOMLINSON AND J. R. VINCENT ......................... 191 The Treatment of Aublet’s Generic Names by His Contemporaries and by Present-Day Taxonomists. PANES Le ZARUCCHD 445 odie dc ovd dawdids dees ven ddewesencds 215 Vegetative Anatomy and the Taxonomic Status of //ex collina and Nemopanthus (Aquifoliaceae). PIRCERCUOBAS. 1.5 649246500 65st neve aes Onn pack cadena ed ones 243 The Chromosomes and Relationships of Metasequoia and Sequoia (Taxodiaceae): an Update. S. E. SCHLARBAUM, T. TSUCHIYA, AND L. C. JOHNSON .......... key Volume 65, Number |, including pages 1-148, was issued January I1, 1984. JOURNAL oF tre ARNOLD ARBORETUM HARVARD UNIVERSITY VOLUME 65 NUMBER 3 US ISSN 0004-2625 Journal of the Arnold Arboretum Published quarterly in January, April, July, and October by the Arnold Arboretum, Harvard University. Subscription price $50.00 per year, plus $5.00 postage for addresses outside of U, 8A Subscriptions and remittances should be sent to Journal of the Arnold Arboretum, P. O. Box 368, Lawrence, Kansas 66044, U.S. A. Claims will not be accepted after six months from the date of issue. POSTMASTER: send address changes to Journal of the Arnold Arboretum, P. O. Box 368, Lawrence, Kansas 66044, U.S. A. Volumes 1-51, reprinted, and some back numbers of volumes 52-56 are available from the Kraus Reprint Corporation, Route 100, Millwood, New York 10546, U.S.A EDITORIAL COMMITTEE S. A. Spongberg, Editor E. B. Schmidt, Managing Editor P. S. Ashton K. S. Bawa P. F. Stevens C, E, Wood, Jr. Printed at Allen Press, Inc., Lawrence, Kansas COVER: The stylized design appearing on the Journal and the offprints was drawn by Karen Stoutsenberger Second-class postage paid at Lawrence, Kansas. BERNICE G. SCHUBERT (photograph taken in 1974). It 1s with respect and affection that the staff of the Arnold Arboretum ded- icates this number of the Journal of the Arnold Arboretum to Bernice G. Schu- bert, Curator of the Arnold Arboretum and Senior Lecturer on Biology in the Faculty of Arts and Sciences of Harvard University. Dr. Schubert retired fromm the staff of the Arnold Arboretum at the end of June, following her seventieth birthday. We are joined by her many friends and colleagues both here at Harvard and around the world 1n wishing her continued good health and many happy years ahead. Dr. Schubert came to the Gray Herbarium of Harvard University in 1935 to pursue her graduate education, and she received the Ph.D. degree from Radcliffe College in 1941. Her thesis research centered on the leguminous genus Desmodium, and the genus has remained one of her primary research interests to the present day. As a result of her studies, she has contributed treatments of Desmodium for numerous regional floras of the United States, Mexico, Central and South America, Africa, and Asia. During the early 1940’s, her interest was also aroused in the New World species of Begonia, and several of her important contributions to the taxonomy of the genus have been completed in collaboration with her friend and colleague, Lyman Smith. After the completion of her graduate training, Dr. Schubert remained at the Gray Herbarium until 1950 as assistant to the Director, Professor M. L. Fer- nald, who was then completing the eighth edition of Gray’s Manual of Botany. For this work Dr. Schubert traveled to Europe in 1946-47 to photograph type specimens in several European herbaria. Her photographs appear in many of Professor Fernald’s papers published in Rhodora and the Contributions from the Gray Herbarium and also in the ten papers they published together. In the final stages of production of the eighth edition of Gray’s Manual, Dr. Schubert saw the book through publication and was, in fact, Fernald’s ‘“‘eyes” for this critical job. Fernald wrote in his acknowledgments, ‘‘Finally, especially since the impairment of his eyesight, the author has had the most loyal, conscientious and unlimited aid of Dr. Schubert in the exacting details of coordinating usages in the text, editing the manuscript for the printer, checking and double-checking the citations of figures and the scores of other details necessary in the book as it goes to press. My appreciation ot all her helpfulness can not be adequately expressed. Cambridge, June 1 In 1950 Dr. Schubert was awarded a Guggenheim Fellowship for study in Europe, and during 1951 and 1952 she remained in Belgium, where she worked on a cooperative project involving the floras of Africa, especially that of the Belgian Congo (Zaire). She returned to the United States in 1952 to work with the New Crops Research Branch of the U. S. Department of Agriculture. It was during this period that she initiated her studies of the taxonomically com- plex genus Dioscorea, the true yams and the group that completes the triad of genera for which she is well known. It was also while she was at the U.S.D.A. that she published with the late J. J. Willaman the fundamental reference, Alkaloid- Bearing Plants and Their Contained Alkaloids. Bernice rejoined the Harvard botanical community in 1962 when she re- turned to Cambridge as a curator of the Arnold Arboretum and as editor of the Journal of the Arnold Arboretum. She filled the latter role from 1963 to 1979, maintaining the Journal as one of the best-edited botanical periodicals of its kind. In this work her kind consideration for authors was unusual, and many of the contributing authors during her term as editor are only too well aware of the debt they owe to her for the improvement of their manuscripts. She was particularly generous in the time and work she was willing to expend on manuscripts received from graduate students, from authors with newly won Ph.D.’s, and from foreign botanists whose English required polishing. In 1979 she assumed chairmanship of the newly established Arnold Arboretum Pub- lications Committee, a post she held until her retirement. Dr. Schubert served as supervisor of the Harvard University Herbaria build- ing from 1969 through 1975, an often thankless position requiring constant attention to the maintenance and day-to-day operation of the herbaria building. She was also appointed Senior Lecturer in the Department of Biology and in this capacity advised numerous undergraduate biology majors and worked closely with graduate students in botany. Over the years Bernice has been unfailing in her encouragement of young botanists; she has helped countless correspondents, including many from Mexico and Central and South America, and numerous amateurs with a deep interest in the genus Begonia. Throughout her career Dr. Schubert has visited Mexico and Central and South American countries to collect specimens for her studies and to participate in botanical activities at several universities and research stations. She has also been a regular participant in Mexican botanical congresses, and she has been named honorary vice president of several of these congresses in recognition of her contributions to Mexican botany and her encouragement and help to young botanists. She was further honored with a vice presidency of the III Congreso Latinoamericano y II Nacional (Peruano) de Botanica in Lima in 1982. We expect that she may wish to escape winter weather in Massachusetts through continued visits ‘south of the border,” but we are hopeful that her taxonomic and curatorial work, as well as her editorial advice, will continue at the Harvard University Herbaria. JOURNAL OF THE ARNOLD ARBORETUM VOLUME 65 JuLy 1984 NuMBER 3 A TAXONOMIC REVISION OF THE AMERICAN SPECIES OF AGARISTA (ERICACEAE)!:? WALTER S. JUDD A GENUS of 30 species occurring in both Africa (including Madagascar) and the Americas, Agarista is closely related to several genera in the Andromedeae (Ericaceae) such as Craibiodendron W. W. Smith, Lyonia Nutt., and Pieris D. Don (see Stevens, 1970, 1971; Judd, 1979). In this paper I have reevaluated the limits of the genus and revised the species occurring in the New World; ca. 1900 herbarium specimens have been examined. Usually the African taxa are treated as the genus Agauria, and the American species as a section of the eastern Asian—North American genus Leucothoé D. Don (see Sleumer, 1938, 1959). The interrelationships and confused taxonomic history of these three groups have been studied by Stevens (1970) and Judd (1979) and are reinvestigated here. The American species of Agarista form a very homogeneous and obviously closely related group, and it has been difficult to delimit phylogenetic groupings of species within this section. However, the taxonomic usefulness of many characters employed in species delimitation (e.g., presence of unicellular hairs and multicellular gland-headed hairs, inflorescence structure, leaf shape) has been reevaluated. I have attempted to employ consistent specific concepts, to compare the taxa and interpret their evolutionary relationships, and to develop practical keys for identification. PHYLOGENY AND EVOLUTION GENERIC AND SECTIONAL RELATIONSHIPS The genus Agarista can be divided into two natural and morphologically distinctive groups that are recognized in this treatment as sections. Agarista ‘The third in a series of revisionary studies of genera of the And deae (Eri ). Previously studied genera include Lyonia (Judd, 1981) and Pieris (Judd, 1982). 2Florida Agriculture Experiment Station Journal Series no. 4262. © President and Fellows of Harvard College, 1984. Journal of the Arnold Arboretum 65: 255-342. July, 1984. 256 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 sect. AGAURIA includes the single widespread and variable species A. salicifolia, which is native to central Africa and Madagascar (see Sleumer, 1938). This taxon is characterized by twigs with nonchambered, slightly to very hetero- geneous pith; leaves with the abaxial epidermis papillose and the adaxial epi- dermis having more or less short, usually undivided cells (in cross section); and capsules with basal placentae. In addition, the style is apparently not swollen. Agarista sect. AGARISTA includes 29 species (34 taxa) and is most diverse in South America, especially southeastern Brazil. These species are characterized by twigs with non- to clearly chambered, Ca//una-type pith (i.e., cells small and thick walled toward the outside of the twig, becoming much larger and thinner in the center); leaves with the abaxial epidermis nonpapillose and the adaxial epidermis with usually tall and often divided cells; and capsules with subapical to more or less central placentae. The style is usually swollen toward the apex. Stevens (1970) has pointed out that the African and American taxa also differ in the development of a hypodermis. Agarista salicifolia has a more or less continuous hypodermis, while the American species have at most one or two cells (in cross section) near the larger veins. Additionally, Cox (1948) has reported differences in the anatomy of the xylem. The sections are very similar in most features. Both are characterized by buds with more than two bud scales; an indumentum of multicellular, mul- tiseriate-stalked, small-headed glandular hairs; leaves usually revolute in bud, with a unifacial midrib bundle and a rather dense vein reticulum in which all orders are more or less equally prominent (Lems, 1964); inflorescences race- mose (or paniculate), overwintering within the bud (Lems, 1962); flowers five- merous, with imbricate calyx lobes; stamens lacking appendages, the filaments geniculate (1.e., S-shaped) and with usually long unicellular hairs; and capsules with nonthickened sutures. Traditionally, Agarista salicifolia has almost always been included in the genus Agauria (see TABLE |) and the American species of Agarista in Leucothoé sect. AGASTIA (see TABLE |; Sleumer, 1959). However, these two groups are actually very similar phenetically and cladistically and are obviously closely related; neither is close to Leucothoé sensu stricto (TABLE 2; Lems, 1964; Stevens, 1970; Judd, 1979). All the characters by which these two sections differ (e.g., pith type, presence of papillae on the abaxial leaf epidermis, placenta position, and presence of a leaf hypodermis) are variable infragenerically —and also sometimes infraspecifically—in at least one closely related genus (Pieris, Lyonia, or Craibiodendron). In addition, the degree of morphological/anatom- ical divergence between Agarista salicifolia and the American species of Agar- ista is comparable to that separating sections within other genera of the An- dromedeae (see TABLE 2; Judd, 1969, fig. 7), while their differences with Leucothoé are of “‘generic magnitude” (TABLE 2; Judd, 1979). The African and American species also form a monophyletic group (see Judd, 1979, fig. 2) based upon the shared derived characters of densely reticulate-veined leaves that are usually revolute in bud. This group is part of a larger monophyletic group that includes Lyonia, Craibiodendron, and Pieris and is characterized by anomo- cytic stomata, fiber bands in the phloem, usually elongated seed-coat cells, 1984] JUDD, AGARISTA 257 TABLE |. Comparative treatment of Agarista and Leucothoé by various authors. TAXON Agarista sect Agarista sect. Leucothoé AUTHOR Ag Agauria (several sections) Lamarck (1783) Andromeda Andromeda Andromeda Michaux (1803), Andromeda Andromeda Nuttall (1818) G. Don (1834) Agarista (but Agarista Leucothoé A, populifolia laced in Leucothoé) De Candolle Leucothoé sect. Leucothoé sect. | Leucothoé sect (1839) Agastia (but Agauria uleucothoé (but 2 spp. placed in 1 sp. placed in Amechania, 21 Zenobia) Leucothoé sect Agauria, 11 Leucothoé sect uleu a Bentham & Hooker Agarista Agauria Leucothoé (1876) sella es a (2 sections) in Leucothoé sect Euleucothoé Gray (1878) Leucothoé Agauria Leucothoé (2 subgenera treated) Niedenzu (1889) Agarista (2 sections, Agauria Leucothoé A, populi olia (2 sections) placed in Leucothoé sect. Euleucothoé) Drude (1897) Leucothoé subg. Agauria Leucothoé subg. Agarista _ A, Euleucothoé and seal ced Lyonia subg ae Enero Eubotrys Small (1914) poe only — Eubotrys, fen hie ser Leucothoé Oreocallis ut E Asia spp. not treated) Sleumer (1938, Leucothoé sect. Agauria Leucothoé 1959) Agastia (6 sections) Agarista Agauria Leucothoé Stevens (1970, 1971) geniculate and/or spurred filaments, and tendency toward epidermal lignifi- cation. The various generic and infrageneric classifications of the species here con- sidered in Agarista are summarized in Tas_e | (see also Stevens, 1970; Judd, 1979). The generic delimitation adopted here is essentially that of G. Don (1834). Lamarck (1783) described—as species of the then very broadly defined 258 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Tas_e 2. Variation in selected pa iat sg anatomical characters in Agarista Leucotho TAXON Agarista Agarista CHARACTER Agarista Agauria Leucothoé Wood Phloem with bands of fibers + + = Pith Calluna-type Slightly to very Variable (often septate) heterogeneous Plant ea + + +/— rie in bud +(—) + — Margin serrate —(+) - + Vein reticulum dense + + — eaten: lignified + + Epidermal c cells divided +(—) —(+) — Abaxial epidermis oe _ + 7 Stomata anomocyti + + —- Anther Fila ave geniculat + + = Filaments with apical + + +/— hair Awns present — = au ae Placenta position Apical (central) Basal Apical Testa cells Elongated Elongated + Isodiametric 1 8 Number of species enus Andromeda—the first known species of both sections of Agarista: A. populifolia of sect. AGARIsTA, and A. salicifolia of sect. AGAURIA. Important nineteenth-century treatments of the group include those of G. Don (1834), De Candolle (1839), Meissner (1863), and Bentham and Hooker (1876). More recently, Sleumer (1938, 1959) has extensively studied both the African and the American taxa, and Kinoshita-Gouvéa (1980) has included the Brazilian species as part of her floristic study of the Ericaceae of Brazil. Stevens (1970) was the first contemporary botanist to reaffirm the close relationship between the African and American agaristas. CLADISTIC ANALYSIS The phylogenetic relationships of the species of Agarista sect. AGARISTA were investigated using the method of Wagner (1961, 1962, 1969, 1980). Twenty- seven characters were used and assigned primitive and advanced states. Most of these characters are quantitative, and the states of such characters are nec- essarily somewhat arbitrarily defined. However, the dividing point of character states was chosen in such a way as to minimize the number of taxa showing 1984] JUDD, AGARISTA 29 WA FiGure 1. Wagner tree for species and varieties of Agarista sect. AGARISTA: extant taxa (black circles); hypothetical ancestors showing only derived character states shared by taxa positioned distally on tree (open circles). Taxa indicated by numerals assigned group, VII = 4. albiflora group, VII = A. revoluta group. (See TaBLe 4 for character states for each species or variety, and for distinctive features of each species grou both conditions. TABLE 3 lists these characters. For each of them, a taxon was scored 0 if primitive and | if advanced. When two or more states of a given character were considered derived, each was given a lower-case alphabetic superscript (e.g., 1°, 1°). TABLE 4 shows all taxa and their character-state values. For each taxon the total divergence index was determined by adding the in- dividual character-state values. Then mutual groupings of derived characters were determined (by hand), and the taxa were arranged in sequence according to these groupings. In the process, an attempt was made to minimize the number of character-state reversals and parallelisms. The taxa were then plotted graph- ically (FIGURE 1), with the branching points determined by the mutual grouping of derived characters and the distance by the divergence of each taxon. The Wagner Groundplan Divergence method, like other cladistic methods, 260 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 TABLE 3. Characters used in the phylogenetic ean of the species and varieties of Agarista sect. Agar CopDE CHARACTER PRIMITIVE STATE ADVANCED STATE A Habit + Well-branched Subshrubs with rigidly shrubs or trees ascending branches B Gland-headed hairs At least sometimes Lacking resent C Stem glaucousness Lacking At least sometimes present D Pith (degree of Nonseptate to At least sometimes septation) regularly ee clearly septate E Pith (cellular Homogeneous Calluna-type structure) F Leaf size Usually > 3 cm Usually < 2.5 cm G Leaf shape Usually + ovate a. Often oblong to elliptic b. + Linear c. Often + orbicular H Leaf apex Acute to rounded- a. Often acuminate mucronate b. Consistently rounded- to retuse- mucronate I Leaf base Not cordate At least sometimes cordate J Abaxial laminar + Lacking or in- At least sometimes lands conspicuous conspicuous K Unicellular hairs on Lacking to sparse Usually + dense abaxial leaf surface 1 Lamina + Flat to ve a. Slightly to strongly slightly revolute — folded at margin . Often oth strongly ene argin c. Often clearly curved toward abaxial surface M Lamina texture Coriaceous, + Very coriaceous, + flexible when dry inflexible when dry N Leaf margin Entire (mature a. Often crisped/undulate leaves) b. Often serrate O Petiole length Short to moderate Often elongate Pp Inflorescence position Axillary Axillary or sometimes termina Q Inflorescence type Raceme a. Raceme or panicle b. Sometimes flowers solitary R Inflorescence length > 3cm <3cm S Inflorescence indu- Hairs (sho a. Nearly always lacking mentum (unicellular) whitish) often b. Hairs ferrugineous present c. Hairs long, whitish T Calyx lobe length Usually < 3 mm Usually > 3 mm U Corolla color White, to pink Often red tinged at apex 1984] JUDD, AGARISTA 261 TABLE 3 (continued). CopE CHARACTER PRIMITIVE STATE ADVANCED STATE Vv Corolla size < 13 mm Often > 13 mm W Corolla pubescence Lacking (or very Sparse to dense (abaxial surface) sparse) x Ovary pubescence At least a few hairs Lacking (unicellular) Y Placenta position + Subapical + Central Z Capsule wall Not stout Stout AA Seed length Usually = 2 mm Usually < 1.5 mm requires numerous assumptions concerning which is the derived and which the ancestral state of each character. The problems and sources of error as- sociated with each step in the procedure are discussed in Judd (1979), Stevens (1980, 1981), Crisci and Stuessy (1980), Wiley (1981), Watrous and Wheeler (1981), and Wheeler (1981). Although the consensus is that out-group analysis 1S mae most powerful method for determining polarity, Crisci and Stuessy main- n that in-group analysis, if used with care, can be helpful in phylogenetic eerena The derived state(s) of characters A-G, I-L, N-P, T, U, W, and ere determined by out-group comparison (using the genera Craibiodendron and Lyoniaas the outgroup; see Judd, 1979), supported by in-group comparison for the character states A, C, F, G, J-L, N-P, T, U, W, and Y. Characters AA and Y are highly correlated. In-group comparison was used in determining the advanced state(s) of H, M, Q-S, V, X, and Z. These decisions were made after careful study of the genus and related groups. The Wagner Tree resulting from the above procedure (FiGuRE 1; see also TABLE 4) indicates that the species of Agarista sect. AGARISTA may belong to eight clearly to rather poorly defined phyletic groups: the Agarista eucalyptoides group (species 25-29), the A. hispidula group (species 17-22), the A. oleifolia group (species 13-16), the A. coriifolia group (species 9-12), the 4. niederleinii group (species 23, 24), the 4. populifolia group (species 6-8), the A. albiflora group (species 1-3), and the 4. revoluta group (species 4, 5). It is also evident that the most advanced taxa—Agarista coriifolia, A. angustissima, A. glaber- rima, A. virgata, A. hispidula, and A. ericoides—are all indigenous to southern Brazil. Most are, in fact, endemic or nearly endemic to Minas Gerais. In contrast, the more primitive taxa—Agarista albiflora, A. subcordata, and A. duckei—are found in northern South America, the two former in the northern Andes and the latter in the Guayana Highland. The species of the Agarista eucalyptoides group are characterized by their distinctive leaves with often elongate, flexuous petioles (O) and acuminate apices (H?). These species also have leaves with rounded to narrowly cuneate bases, white flowers with short calyx lobes, and capsules with subapical pla- centae. Agarista eucalyptoides and A. boliviensis are likely closely related. Both taxa lack gland-headed hairs (B), have often crisped/undulate leaf margins (N°), and have leaves that are revolute in bud and more or less flat at maturity. In contrast, the more advanced A. duartei, A. angustissima, and A. glaberrima 262 JOURNAL OF THE ARNOLD ARBORETUM TaBLe 4. Character divergence [VOL. 65 values for taxa used to construct Wagner Tree. eae BCDEFGHIJKLMNOPQRSTUVWXYZAA~ Geog. Total Taxon ] o0001T000N/0000000001cCO0000000 A 3 2 010017100Nj/000000000000000007? A 4 -.3_...091001 100/000 bOD0000000001000 AL _ 5. _ 4 0j1j0o01000000 ido o0000000000000 4G 3 5a 0j1;0 0 100 1b00 0 ido 00000000000000 B-N 4 __ bb. _ oftfo.01 00 1D001|1¢009000000010077 BN | 6. 6a 0]1/O}1]}1 0 0 140 o0000000/1}000010000 NA 6 6b OjJ1JO}1]1 0 0 ido 01000000j]1}/000010000 NA 7 7 0}1}O/1}1 0 0 ido 00000000]1}000001000 44NA 6 8. ofofolili o ofiza0000 moc ohloogg01900 mA 6 | 9a 011/71 11000141J0 0ONjJo011a0001000000_ B-N 10 9b 0717/1 110 1a0 0]1/0 Of1J0011a0001000000 B-N 10 10 O11j0011001;/1J00N)/0o0000000000001 B-N 7 1] 0101/00 1100141/1 0fF1}000001c0101000 0? B-N 9 _ 12 _ _ of.}0.0101a0 Ofof1 olofoooVO1cOO010000 BN 6 | 13 00001000]1/0100000000 Oj0/0 1 OF140)1 B-S 6 14a 00011000/1]/0000000000 0f1}0 0 0}1JO}1} B-N 6 14b 00011000/0}000000000 la0}1}1 01)4140}1 B-N 8 15 0101101 0/0/000000001 0 OJ0J0 0 0/1)0]1} B-S 7 16a 0000100 0/1}0000001 1a0 0 0f1/0 0 0}1}0}1 B-S 7 __ lb _ 000011 00]1Jo0000000000}1]0 0 olololl}. BN _ 5. _ 17 1/0 00 1]1Jlclb1jO OJFOJO000000/1}/0000000 B-S 7 18 1/0 0014/1/0 0/1/0 OJOJO 0071 1a0 1;O}0001101 B-N 10 19 1/0 0 0141/0 0/1/00 0/16000100011/0000000- B-NS 7 20 0?1 01 1}1}0 OJ1J/0 O}1bO 000 1b1 OJ/OJ0000000- B-N 8 21 1/0 001/1)0 0/1}/0 0J/160000000]1/1010101 B-N 10 _ _ 22, _ _I]o.0.0 141}0 ofi}o ofpo 90001 01/2 000101_ BN. _ 10. _ 23a Oy1yON}1 171000000000 O0/O)0000000)1]0 B-S 6 23b O;1;01141 017 1aADO OAOD0G0A0O0;1j0 00000 0)1j0- B-S 7 _. 24. _ Ofllolz]2 001020000000 0f1Jo000000]110_ BN 6 | 25 010010 0}/la0 0 0/0J0 laljoo1000001000 A 7 26 01001 0 1ala0 0 0/0}0 1aljO001b00000000_- B-NS 7 27 010110 O}la0 0 O}1a0 0J1]1 1aA0 1aA00001000— B-N 10 28 00001 0 lblaO 0 O}1a0 0}1}1 1aA0 1800001000 B-N 9 29 10001 0 Ibla0 0 Of]a0 OJ}0/001000000000— B-N 6 Explanation of abbreviations: A = Andes rth Highland, NA = N , B = Brazilian region (north and south), G = Gua closed within de markings. Each taxon indicated by its species number. aya merica. Dotted aes separate species groups supported by characters en- na 1984] JUDD, AGARISTA 263 have leaf blades that are slightly to strongly adaxially folded. The distinctive linear leaves (G*) of A. angustissima and A. duartei have probably evolved convergently. Except for A. boliviensis, which occurs in the southern Andes, all species of this phyletic group are limited to the Brazilian region (see geo- graphic analysis). The group is most diverse in Minas Gerais, with only the relatively widespread 4. euca/yptoides extending southward into the state of Rio Grande do Sul and adjacent Uruguay. The species of the Agarista hispidula group are characterized by their low, more or less sparsely and erectly branched habit (A) and their small, usually cordate-based leaves (F, I). Many species also have strongly revolute leaves (L») and elongate calyx lobes (T). The position of the placenta varies from more or less subapical to central, and the group may be derived from extinct, small- leaved members of the 4. o/eifolia group, an assemblage with mainly central placentae (note especially the resemblance in leaf characters to A. pulchella var. cordifolia). Although A. hispidula and A. ericoides are probably closely related, both often having red corollas, more or less centrally positioned pla- centae, and short seeds, the relationships between these species and A. chlor- antha, A. organensis, A. nummularia, and A. virgata are somewhat obscure. Species of the A. hispidula group are limited to the Brazilian region. The group is most diverse in the northern subunit of this region, with only A. chlorantha and A. nummularia occurring as far south as Santa Catarina (the former) or Rio Grande do Sul (the latter). The Agarista oleifolia group is a rather loosely knit assemblage of taxa char- acterized by often cordate-based leaves (I), and by capsules with central pla- centae and short seeds (Y, AA). These species tend to have more or less flat leaves and flowers with short calyx lobes. Several taxa have straight-sided or oblong leaves (G*). The relationships between the taxa comprising this rather weakly defined group are obscure. The group is limited to the Brazilian region (including Paraguay). The four species of the Agarista coriifolia group are characterized by their lack of multicellular gland-headed hairs (B), and by their usually quite coria- ceous leaves (M) with often conspicuous abaxial glandular regions along the midvein (J). The leaves tend to be more or less flat, the inflorescences elongate, and the placentae subapical. The calyx lobes are usually short, and the flowers tend to be reddish. The affinities of A. chapadensis are probably with A. sub- rotunda, a species with moderately to consely pubescent abaxial leaf and corolla surfaces (K, W). The species of this y line are limited to the northern portion of the Brazilian region (see phytogcosraphic discussion). The Agarista niederleinii group includes only three taxa, which are distin- guished by their lack of multicellular gland-headed hairs (B), their often septate pith (D), and their large capsules with very stout walls (Z). These taxa also tend to have short racemes (R), subapically positioned placentae, white flowers with short calyx lobes, and flat to slightly revolute leaves. The group is limited to the Brazilian region. The Agarista populifolia group is distinctive due to its often clearly septate 264 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 pith (D), its ovate, acuminate leaves (H*), and its short racemes (R). The members of this lineage frequently have rather large, more or less flat leaf blades with cuneate to rounded bases, white flowers with short calyx lobes, and capsules with subapical placentae. These species are limited to North and Central America, growing in the southeastern United States and east-central Mexico south to Honduras and El Salvador. This group may have originated from extinct members similar to species of the A. al/biflora group (note especially the resemblance of leaves to those of A. albiflora). All three species re g the Agarista albiflora group occur in the northern Andes. These species are likely rather primitive, and although it is clear from morphological and geographic evidence that the small-leaved 4. bracamorensis and A. subcordata were probably derived from ancestors similar to the variable and widespread 4A. albiflora, these three species are only weakly linked in at least sometimes having cordate leaves (1). Finally, the two species comprising the Agarista revoluta group (VIII) are characterized by their lack of gland-headed hairs (B; except rarely present on the ovary of A. duckei!) and their leaves with the blade often quite variable in extent of abaxial curvature (L‘). The capsules are often ovoid, with the valve margins sometimes slightly differentiated; the placentae are subapical, and the seeds are the longest (2—-3.5 mm) of the section. The primitive species, 4. duckei, occurs chiefly in the Guayana region, while the slightly more specialized A. revoluta grows in coastal “‘restingas” from the state of Bahia south to Estado do Rio in eastern Brazil. There are insufficient data to determine the evolutionary interrelationships of these eight cladistic groups; all share the derived character-state of Ca/luna- type pith (E). In contrast, the African species, Agarista salicifolia, has slightly to strongly heterogeneous pith (Stevens, 1970). The species of Agarista sect. AGARISTA are obviously all closely related, although the extensive parallel evolution of characters and the frequency of character-state reversals obscure both intra- and intergroup relationships. Many examples of parallel evolution can be found among the various phylogenetic groups discussed above; these can be determined from FiGure | and TABLE 4. Of the 27 characters used, at least 19 show some degree of intergroup parallelism (and also often intragroup parallelism or reversals), while 2 (O, T) of the remaining 8 show intragroup reversals and | (E) is invariant. In groups showing a large amount of homoplasy, such as Agarista sect. AGARISTA, the elucidation of cladistic relationships be- comes extremely difficult. Thus, the conclusions reached here should be inter- preted as very preliminary hypotheses, to be tested by the evaluation of ad- ditional characters (including anatomy and chemistry). DISTRIBUTION AND ECOLOGY As discussed above, the 30 species of Agarista recognized here are divided into two sections. Section AGAURIA, including the single species A. salicifolia, occurs in central Africa, Madagascar, Réunion, and Mauritius (Sleumer, 1938), while section AGARISTA, containing the remaining 29 species, occurs mainly in South America (Map 1). Both sections are composed of basically montane 1984] JUDD, AGARISTA 265 L710 104 oO 1000km O 600 mi 304 30 100 Map |. General distribution of Agarista sect. AGARISTA, showing center of diversity in southeastern Brazil. Pattern indicates number of taxa present: stipples, 1; vertical lines, 2; horizontal lines, 3 to 8; solid black, region of greatest diversity. plants, with A. salicifolia occurring between 700 and 3500 m alt. and the many South American species growing from ca. 500 to 2700 m alt. However, A. duckei, a species of the Guayana Highland, can be found as low as 100 m, and A. revoluta, a taxon of coastal “‘restingas” in Brazil, frequently occurs near sea level. (Agarista nummularia and A. pulchella have also occasionally been collected at very low elevations near the coast.) Possible origins of this inter- esting montane tropical transatlantic distribution pattern shown by Agarista are discussed by Stevens (1970). Section AGAURIA occurs in several widely scattered montane areas of central Africa but reaches its greatest morphological diversity in Madagascar (Sleumer, 1938). Species of section AGARISTA occur in five major geographic areas (see TABLE 5): the Coastal Plain of the southeastern United States (1 sp.—Agarista populifolia), the Mexican region— mountainous areas from central Mexico south to Honduras (2 spp.—A. mexicana and A. sleumeri), the Guayana Highland (1 sp.—A. duckei), the Andean region—mountainous areas from Bolivia to 266 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 TABLE 5. Distribution of taxa of patie cies the nations* of North, Central, and South Amer 1. Unitep States (1) . duarteit A. populifoliat A. ericoidest 2. Mexico (3 A. eucalyptoides A, mexicana var. mexican A. glaberrimat . mexicana var. ean A. hispidu A. sleumert A. oleifolia var. glabra 3. GUATEMALA (1) A. oleifolia var. oleifolia A. mexicana var. mexicana A, pulchella var. cordifolia 4. BELIZE 2 A. pulchrat A. mexicana var. mexicana A, revoluta var. revoluta nexicana var. pinetorum A, subrotundat 5. EL pein A. virgatat A. mexicana var. pinetorum g. Estado do Rio/Guanabara (7) 6. HonpurRas (2) A, Se var. eens A, mexicana var. A. eucalypto A. mexicana var. a A. hispidu le 7. COLomBIA (1 A. oleifolia var. oleifolia albiflor A. organensi 8. Ecuapor (1) A, revoluta var. revoluta A. albiflora ean 9. Peru (3) h. SAo PAULO (6) A. albifl chlorantha A. bracamorensist A. eucalyptoides subcordata A. hispidul 10. VENEZUELA (2) yi oleifolia var. oleifolia . albiflora pulchella var. cordifolia A, duckei a Soe var. pulchella 11. Botrvia (1) i. Parana eh A. boliviensist A. chlor 12. Brazit (25) A, ey var. acutifolia Para (1) A. niederleinii var. niederleinii duckei A. pulchella var. pulchella b. Bahia (5) j. Santa Catarina (7) A. chapaden A. chlorantha A coriifolia v var. coriifolia A. eucalyptoides A. oleifolia var. glabra A, minensis A. revoluta var. revoluta A. niederleinii var. acutifolia A. revoluta var. velutinat A. sseadlae var. niederleinii c. Goias (2) A, numm A. chapaden A, uh var. di A. oleifolia var. glabra k. Rio Grande do Sul (5) d. Dist. Federal A. eucalypto oides chlorantha A, sed var. glabra ; niederleinti var. acutifolia e. oe osso (2 A. niederleinii var. niederleinii el A. nummularia oleflia var. glabra 13. Urucuay (1 f. wee Gerais (16) A. eucalyptoides A, ae ete 14. PARAGUAY A. chlorantha araguayensis A. cortifolia var. coriifolia 15. ARGENTINA (1 A. corlifolia var. bradeit A. paraguayensis *Within Brazil, states. +Taxon endemic to nation or state. 1984] JUDD, AGARISTA 267 Colombia (4 spp.—A. albiflora, A. subcordata, A. bracamorensis, A. bolivien- sis); and the Brazilian region— mountainous areas of southeastern Brazil (and adjacent Uruguay and Argentina) (22 spp.). With the exception of 4. duckei, which has many populations in the Guayana Highland and a disjunct locality in western Mato Grosso, all species are endemic to one of these five regions. The extremely diverse Brazilian region can be divided into northern and southern subunits, with the northern including the mountains of Minas Gerais and the adjacent states of Sdo Paulo (northern part), Rio de Janeiro, Guanabara, Bahia, Goids, and Mato Grosso, and the southern mountainous regions of the states of Sdo Paulo (southern part) and Paranda south to Rio Grande do Sul and adjacent areas of Uruguay, northeastern Argentina, and Paraguay. The northern subunit of the Brazilian region has the most species, with eighteen that are indigenous (see Map 1). Within this area, the state of Minas Gerais has by far the most species, with fourteen (sixteen taxa), of which seven (eight taxa) are endemic; Estado do Rio/Guanabara is next with seven indig- enous species (two endemics), followed by Sdo Paulo (northern) with five species (no endemics) and Bahia with four species (one endemic variety). The southern subunit contains only seven indigenous species (eight taxa), of which four species (six taxa) are endemic. Only three species, Agarista pulchella, A. chlor- antha, and A. eucalyptoides, occur in both subunits. TABLE 5 lists the species and varieties of Agarista occurring in or endemic to the various countries (and within Brazil, states) of North and South America. The mountainous region of southeastern Brazil is a major center of diversity for many groups of vascular plants (see L. B. Smith, 1962; Good, 1964; Tryon, 1972). Agarista appears to have undergone extensive speciation (probably eco- geographic) in these mountains, since this region supports several groups of very derived taxa such as A. coriifolia, A. glaberrima, A. angustissima, A. virgata, A. hispidula, and A. ericoides. This concentration of Agarista species may be a relatively recent development, formed in response to Tertiary climatic and geologic changes (see Stevens, 1970). The area of origin of the group is unknown. It is of interest that the primitive species Agarista duckei occurs in the floristically diverse Guayana Highland, while two other rather primitive taxa (A. albiflora and A. subcordata) occur in the geologically recent Andean region. garista populifolia, of the southeastern United States, has its closest affinities with the montane Mexican taxa A. sleumeri and A. mexicana. There are many similarities between the deciduous forest formations of the eastern United Dressler, 1954; Martin & Harrell, 1957; Graham, 1964, 1973; Rzedowski, 1965; and Gémez-Pompa, 1973). A similar disjunction pattern is seen in the related genus Lyonia, with L. ferruginea (Walter) Nutt. and L. fruticosa (Mi- chaux) G. Torrey occurring in the southeastern U. S. and L. squamulosa Mar- tens & Galeotti in the Sierra Madre Oriental of Mexico (Judd, 1981). Although a few species such as Agarista oleifolia, A. albiflora, A. mexicana, and A. eucalyptoides have quite wide distributions, most (e.g., 4. pulchra, A. virgata, A. organensis, A. ericoides, A. angustissima, A. duartet, A. bracamo- 268 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 rensis) are narrowly endemic. Most varieties and closely related species are more or less geographically isolated, but a few taxa, such as the varieties of A. niederleinii, seem to be chiefly elevationally/ecologically isolated. e genus grows in a wide variety of habitats, but most species prefer acid soils and are sun-loving plants of open communities. Most also sprout readily from the base after fire or disturbance. Characteristic habitats include cerrado, rocky or grassy campo, boggy or marshy campo, shrub bogs, gallery forests along rivers or streams, thickets, open scrub, forest or thicket margins, elfin woodland or moist montane forests, rocky crests or slopes, coastal scrub/ “‘restinga,” sand dunes, savannas, and pine-oak forests. For detailed discussion of cerrado vegetation (along with lists of characteristic species), see Eiten (1972). Species may occur in iron-rich lateritic soils, sandstone regions, white sands, or highly organic soils. TAXONOMIC CRITERIA Because of the extensive parallel evolution and character-stat al in Agarista, different species can be separated only by using rete OF ee and reproductive characters. Morphological entities that have no major internal discontinuities and that are separated from other, similar entities by consistent morphological gaps are considered to be species. They have definite geographic distributions and ecological preferences. Many species, such as 4. pulchra, A. virgata, A. organensis, A. ericoides, A. angustissima, A. duartei, and A. bra- camorensis, are narrowly endemic. Most species (and varieties) are probably isolated by differences in distribution, ecology, and/or genetic composition. The term “‘variety” has been used for the morphologically distinctive geo- graphic (or ecological) subunits comprising widespread and/or variable species (see Judd, 1981). Varieties have only very slight morphological gaps between them, may intergrade, and often differ in only one (or very few) aspect(s) of indumentum or leaf morpholo The taxonomic usefulness of (and the variation within) the various characters used in this revision are discussed below. Characters most useful in species delimitation include: 1) leaf vernation, size, shape, apex and base types, petiole length, and extent to which the margin is revolute; 2) inflorescence length; 3) unicellular indumentum of the abaxial leaf surface and of the inflorescence axes and other reproductive structures (especially calyx, corolla, and ovary); 4) calyx length and, to a lesser extent, corolla color and length; 5) placenta position (in fruit); and 6) seed length. Hasir The species of Agarista vary from small, sparsely and erectly branched sub- shrubs such as A. ericoides, A. virgata, A. hispidula, A. duartei, and A. num- mularia to moderate-sized, laxly branched trees up to 6—8(—20) m tall such as A, populifolia, A. sleumeri, and A. mexicana. The branching pattern, including internode length, density of branching, and orientation of branchlets, is quite variable but is sometimes useful in distinguishing between species (e.g., A. 1984] JUDD, AGARISTA 269 virgata and A. pulchra, see key). Although the bark is always longitudinally furrowed, it is very poorly developed in the smaller species (e.g., A. virgata, A. duartei, A. ericoides). It 1s occasionally of taxonomic interest, such as in separating A. populifolia, in which the bark is thin and shallowly furrowed, from the related A. mexicana and A. sleumeri, in which it is thick, corky, and deeply furrowed. Cerrado species such as A. eucalyptoides tend to have con- torted trunks and thick, corky bark, characters very common in this vegetation type (see Eiten, 1972). Most species sprout readily from the base after fire or disturbance. As in most other members of the Andromedeae, the shoot meristem in Agarista has a limited life span, and growth in height is achieved by a succession of equivalent axillary, orthotropic shoots (Lems, 1962). All species show Leeu- wenberg’s Model of growth (see Hallé, Oldeman, & Tomlinson, 1978, for description) or a slight variation of this architectural pattern. Floral buds are usually produced in the leaf axils of the distal portion of the shoot, although they may be restricted to those at or very near the apex. The shoot apex typically aborts, but in some species (e.g., 4. angustissima, A. coriifolia, A. glaberrima) it is converted into a terminal inflorescence (with the leaves gradually grading into bracts). The inflorescences overwinter within the buds, with meiosis pre- sumably occurring in the spring (see Lems, 1962). TwIGs The branchlets vary from terete to slightly angled. Pith is of the Calluna- type in all American species and varies from non- to clearly chambered. Its structure is frequently of taxonomic value (see key) and is especially useful in separating Agarista paraguayensis, A. populifolia, and A. sleumeri from A. mexicana. INDUMENTUM The indumentum of Agarista is of three distinct types: unicellular hairs; multicellular, multiseriate-stalked, gland-headed hairs; and papillae. Although the presence or absence of multicellular, gland-headed hairs has been much used in the taxonomy of the genus (Sleumer, 1959), this character 1s actually too variable to be of much taxonomic significance. Of the 29 species comprising Agarista sect. AGARISTA, 13 have both glandular-pubescent and nonglandular forms. Only A. virgata is consistently glandular-pubescent. However, this taxon has been very poorly collected and nonglandular forms may eventually be discovered. Fifteen species consistently lack glandular hairs. Glandular-pu- bescent and nonglandular individuals frequently occur intermixed in the same population (e.g., Judd 2609, A. populifolia). There is also much variation in density and distribution of multicellular gland-headed hairs. At one extreme are plants with glandular hairs on the young twigs, midvein (or even lamina) of both leaf surfaces, inflorescence axes, pedicels, and calyx lobes; other indi- viduals may have such hairs only on the inflorescence axes, or only along the leaf margins (i.e., leaves + glandular-ciliate). In a few species such as A. an- gustissima and A. duartei, glandular hairs are only present on juvenile plants. 270 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 65 Finally, A. duckei is unique because of its occasionally glandular-pubescent capsules. However, because of the extensive intrapopulational (and develop- mental) variation in glandular indumentum, it has not been possible to use this character at either the specific or the infraspecific level. The several groups of extremely similar taxa (e.g., A. nummularia var. hummularia and Leucothoé (Agarista) nummularia var. floccigera, A. chlorantha and A. serrulata; A. his- pidula and A. intermedia, A. ericoides, Leucothoé (Agarista) lycopodioides, and L. (Agarista) acicularis, A. subrotunda and A. pistrix; and A. oleifolia var. oleifolia and Leucothoé (Agarista) oleifolia var. hispidula) that were recognized by Sleumer (1959) and that differ chiefly in the presence or absence of such hairs have been considered conspecific. All species of Agarista have at least a few unicellular hairs, and the distri- bution and density of such hairs is frequently of taxonomic significance (see keys). Several taxa—A. mexicana var. pinetorum, A. revoluta var. velutina, A. indumentum of the inflorescence axis and other reproductive structures (es- pecially calyx, corolla, and ovary) was found often to be of systematic value. Thus, the inflorescence indumentum is useful in distinguishing 4. glaberrima from A. boliviensis and A. eucalyptoides, A. virgata from A. nummularia, A. oleifolia var. oleifolia from var. glabra, and A. pulchra from A. pulchella. Several taxa (e.g., A. paraguayensis, A. revoluta var. velutina, and A. hispidula) are easily distinguished from related ones due to their abaxially pubescent corollas, and the indumentum of the ovary and calyx is very useful in separating A, mexicana, A. sleumeri, and A. populifolia. However, such characters must be used with care since many species are quite variable in indumentum. Al- though the unicellular indumentum of the twigs 1s usually too variable to be of much taxonomic use, it is distinctive in a few species. For example, the hairs are ferrugineous in A. eucalyptoides and elongate in A. albiflora. Abaxially papillose leaves are found only in the African species, Agarista salicifolia (sect. AGAURIA); all the American species (sect. AGARISTA) lack such papillae. LEAVES Characters of leaf vernation, size, shape, apex and base types, thickness, petiole length, and extent to which the margin is revolute are extremely im- portant in the taxonomy of Agarista sect. AGARISTA. Most species of Agarista have leaves that are revolute in bud; consequently the mature leaves often have a faint longitudinal “fold” line on each side of the midvein. However, A. duartei, A. angustissima, and sometimes A. glaberrima have strongly adaxially folded leaves. The mature leaves may be more or less flat with a plane to only very slightly revolute margin (many species), strongly to slightly adaxially fold- ed (A. duartei, A. angustissima, A. glaberrima), or clearly abaxially curved with more or less strongly revolute margins (e.g., 4. organensis, A. hispidula, A. chlorantha, A. ericoides). There is great variation in size and shape of the lamina—from 0.4—-1.2 cm long in A. ericoides to 4-13 cm long in A. sleumeri, 1984] JUDD, AGARISTA 271 and from linear in A. angustissima to often nearly orbicular in A. nummularia. Numerous examples of the taxonomic usefulness of leaf shape and size are evident in the key. The petiole ranges from very short (ca. 1.5 mm in A. ericoides) to very long (to ca. 40 mm). The often long, slender, flexuous petioles of A. eucalyptoides, A. glaberrima, and A. boliviensis are especially distinctive. The apex and base of the lamina, although often showing much infraspecific variation, are sometimes useful at the species level. Many species have leaves that vary from acute to rounded with a small mucro at the apex, and from slightly cordate to rounded or cuneate at the base. The leaf apex is useful in distinguishing 4. duckei, A. revoluta, and A. albiflora, the base, in separating A. minensis from A. pulchella (see key). Agarista populifolia, A. mexicana, and A. sleumeri are distinctive due to their consistently ovate leaves with acuminate apices. Finally, although most species have moderately coriaceous leaves, 4. coriifolia and its relatives have thickly coriaceous blades that are rather in- flexible when The leaves of most species-are entire but those of Agarista boliviensis are distinctive in having crisped/undulate margins, and those of A. populifolia are often serrate. In this species each tooth is associated with a multicellular gland- headed hair. The margins of several Brazilian species are sometimes glandular- ciliate due to the presence of multicellular gland-headed h Many species have glandular “‘patches” or “dots” along culayei on the abaxial epidermis. These glandular regions (which should not be confused with the much smaller gland-headed hairs found in some species) seem to be as- sociated with the secondary veins and are variable in size and shape. They are sometimes of taxonomic interest, as in distinguishing the varieties of Agarista coriifolia. All species have reticulodromous venation, with a rather dense vein retic- ulum and all orders more or less equally prominent (Lems, 1964; Judd, 1979). INFLORESCENCE The flowers of Agarista are borne in axillary (or, less commonly, terminal) racemes or panicles on branches of the previous season, although in A. organ- ensis the inflorescences are often reduced to solitary axillary flowers or few- flowered clusters, and in several species (e.g., 4. minensis) the axis is so short- ened that the inflorescence is nearly a fascicle. There is always a single inflo- rescence per leaf axil. Terminal flowers are usually lacking, with the apex of the raceme simply aborting, so the inflorescences are polytelic (Weberling, 1965). However, in A. populifolia a few of the flowers composing the raceme are rarely replaced by two-flowered clusters—i.e., one of the bracteoles of the pedicel subtends a secondary flower. This also occurs in A. pulchella and A. coriifolia var. bradei, resulting in a small panicle in which the two- to many- flowered inflorescence branches have terminal flowers, although the apex of the primary axis aborts. However, the apices of even the secondary axes (branches) of many panicles abort (see panicles of A. coriifolia, A. virgata, A. glaberrima). The few-flowered axillary fascicles (or short racemes) of A. or- ganensis also at least sometimes have terminal flowers, and it is probable that 212 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 the pedicel of the ‘solitary axillary flower” with its several bracteoles actually represents a very reduced inflorescence axis (1.e., a terminal-flowered raceme). hus, monotelic (or partly monotelic) inflorescences occur in a few species of the genus. This inflorescence type is, according to Weberling (1965), more primitive than the polytelic type, and the presence of more or less monotelic inflorescences in this group (along with their variation in position) may indicate its low level ofadvancement. In reproductive characters most species of Agaris- ta are certainly less specialized than those of Lyonia, Craibiodendron, or Pieris (Judd, 1979). There is great interspecific variation in inflorescence length—from at most ca. 1.5 cm in ata organensis or A. minensis to over 20 cm in A. coriifolia. This cl lly useful and has been employed in separating A, minensis ae vi pilchella A. angustissima from A. duartei, and A. organ- ensis from A. chlorantha, among others. FLOWERS All species have actinomorphic, perfect, pendulous flowers, although the pedicel curves upward as the fruits begin to develop, ultimately placing the capsules in an erect position. The flowers of all species are five-merous and quite fragrant. Those of Agarista populifolia are visited chiefly by bumblebees (Bombus), no information is available on the pollination of the many South American species. The pedicel varies in length from only |.5-3 mm in Agarista bracamorensis to 14-15 mm in A. angustissima, A. coriifolia, and A. pulchella. This is not a very useful taxonomic character since the pedicel tends to elongate as the flowers and fruits develop. All species have a clearly developed articulation between the pedicel and the receptacle, its position varying from at to ca. 3 mm below the insertion of the calyx. In all species the pedicel is subtended by a single bract, the size of which is usually too uniform to be of taxonomic importance. Each pedicel in Agarista usually has a pair of bracteoles, although there are occasionally three to many in a few taxa. The bracteoles are usually positioned along the pedicel from the base to near the midpoint; they may rarely even be found near its apex. The bracteoles are characteristically minute (see descrip- tions) and linear to triangular or ovate-triangular; very rarely they subtend secondary flowers. The number, size, and shape of the bracteoles are too uni- form to be useful at the species level. The calyx is composed of five persistent, more or less triangular, imbricate sepals that are connate only at the very base. Stomata are present only on the abaxial surface (Stevens, 1971). The lobes show some interspecific variation in length (e.g., 0.5-1.5 mm in Agarista bracamorensis, 3-6 mm in A. ericoides). This character is thus frequently of taxonomic significance, and it has been used to distinguish A. chlorantha from A. organensis and A. nummularia from A. pulchella. The apex varies from acuminate to acute and is usually too uniform to be of systematic importance. However, 4. ch/orantha typically has acute tips, while those of the related 4. Aispidula are usually acuminate. The corolla is always sympetalous and deciduous. Stomata are present on 1984] JUDD, AGARISTA 213 the outer surface but lacking on the inner (Stevens, 1971). In most species the corolla is more or less cylindrical, but in a few such as Agarista albiflora, A. duckei, and A. hispidula it may be urceolate or nearly so. The corolla is most often white (or greenish white) or white with a pinkish to reddish tinge toward the mouth, but in some species (e.g., 4. revoluta, A. coriifolia var. bradei, A. oleifolia var. glabra, A. ericoides) it is frequently red. Its length varies from 5— 8 mm in A. niederleinii var. acutifolia to 10.5—16 mm in A. oleifolia var. glabra. The shape, size, and color of the corolla are only slightly taxonomically useful in Agarista, due both to the overall uniformity of these characters within the genus and to their considerable variation in some taxa. The flowers of all species of Agarista have ten stamens. These are arranged in two whorls and inserted at the base of the corolla; the outer whorl is opposite the corolla lobes. The stamens are always included and are arranged in a tight ring, with the anther pores facing inward. The filaments are slender, flattened, geniculate d, and slightly swollen near the base. They are covered with long unicellular hairs, especially near the base. Filament length varies from 3—-4.5 mm in A. eucalyptoides and ca. 3.5 mm in A. subcordata to 6.5- 8.5 mm in A. oleifolia var. glabra. A white line or triangular patch of disin- tegration tissue is present on the back of each anther lobe near the apex, but this tissue is only poorly developed in several species. Because of the consistency of the androecial characters within Agarista, they have seldom been used at the species level. The ovary is superior. The placentae are axile and are borne subapically to basally on a central columella. Species with more or less central (e.g., Agarista oleifolia, A. pulchella, A. paraguayensis) or basal (e.g., A. salicifolia) placentae have very deeply impressed styles. Placenta position is an important taxonomic character and frequently links related groups of species—for example, sect. AGAaRISTA (subapical to central) vs. sect. AGAURIA (basal), and the 4. oleifolia species group (+ central) vs. the A. coriifolia group (+ subapical). The ovary shape varies from ovoid to subglobose, but this variation (like that in placenta position) is better expressed in the mature fruit. FRUITS AND SEEDS The fruits of Agarista are five-valved, subglobose or short-ovoid to ovoid, loculicidal capsules with pale, unthickened (or occasionally very slightly thick- ened near apex) sutures. Within Agarista sect. AGARISTA, the capsules tend to be rather uniform in size and shape, but those of A. niederleinii and A. uleana are distinctive due to their large size and their thick walls. Placenta position is best observed 1 in mature fruits and, as mentioned above, is a very valuable taxonomic ter within the genus. The variation in placental position with- in Agarista sect. AGARISTA is shown in TABLE 4 and in the key. The seeds are very small, brown, and scobiform, and they have a thin testa composed of very elongate cells. Species with central placentae tend to have short (0.5-1.4(-1.6) mm) seeds, while those with subapical placentae usually have longer (1—3(-3.5) mm) ones. The seeds of Agarista salicifolia are 3-4 mm long (Sleumer, 1938). 274 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 65 VEGETATIVE ANATOMY The vegetative anatomy of the genus has not been studied in detail. However, pith type was found to be an important taxonomic character: it is Ca/luna- type in all American species (see Stevens, 1970) and slightly to very hetero- geneous 1n the African species, Agarista salicifolia (Stevens, 1970). In addition, it remains more or less solid in many American species but becomes nearly hollow in A. paraguayensis, or irregularly to regularly aie agpreectan et in several taxa (see descriptions). The pith is always chambe n A. populifolia, A. sleumeri, and A. minensis. Stevens (1970) has pointed out i A. salicifolia has epidermal cells that are usually not tall and are rarely divided periclinally, and a single-layered hypodermis that is more or less continuous in its leaves. In contrast, the American species of Agarista have epidermal cells that are usually tall and with periclinal dividing walls, and a hypodermis of at most one or two cells (in cross section) near the larger veins. The vascular bundles of the midrib and petiole are unifacial, with very prominent bundle-associated fibers (Niedenzu, 1889; Stevens, 1970). The xylem anatomy has been studied by Cox (1948), who placed A. salicifolia in his tribe Cassiopeae and retained the species of Agarista sect. AGARISTA within the Andromedeae. TAXONOMIC TREATMENT Agarista D. Don ex G. Don, Gen. Syst. 3: 837. 1834. Leucothoé D. Don sect. Agastia DC. Prodr. 7: 603. 1839. Leucothoé D. Don subg. Agarista (D. Don ex G. Don) Drude in Engler & Prantl, Nat. Pflanzenfam. IV. 1: 42. 1889. LecToTyPE SPECIES: Agarista nummularia (Cham. & Schldl.) G. Leucothoé D. Don sect. Agauria DC. Prodr. 7: 602. 1839. Agauria (DC.) J. D. Hooker in Bentham & Hooker, Gen. Pl. 2: 586. 1876. Lecrorype species: Agauria salicifolia (Comm. ex Lam.) J. D. Hooker (= aeaia salicifolia (Comm. ex Lam.) G. Don). Amechania DC. Prodr. 7: 578. 1839. LEcrotyPe SPECIES: Amechania subcanescens DC. (= Agarista chlorantha (Cham.) G. Don) Evergreen shrubs or trees with longitudinally furrowed bark and terete to slightly angled branches; pith Ca//una-type [or slightly to very heterogeneous], nonchambered to clearly chambered. Indumentum of unicellular hairs and often multicellular, multiseriate, long-stalked, gland-headed hairs. Buds conical or slightly flattened and triangular in outline, with 2 to 4 (to 6) imbricate scales. Leaves alternate (to subopposite or nearly 3-whorled at some nodes), simple, petiolate, revolute (or convolute) in bud, often reddish on young shoots, co- riaceous, frequently with gland-headed hairs on midvein or along margin (rarely also on lamina), often with unicellular hairs on midvein (and densely covering abaxial surface) [sometimes papillose abaxially]; margin entire (or undulate, to obscurely to clearly serrate, or serrulate/ciliate due to presence of gland-headed hairs); venation reticulodromous, the vein reticulum rather dense and with all 3Characters given within brackets apply only to the African species, A. salicifolia. 1984] JUDD, AGARISTA Zdd orders more or less equally prominent; petiole vascular bundle unifacial. In- florescences overwintering within bud with meiosis occurring after emergence, axillary (or terminal) racemes or panicles; flowers perfect, 5-merous, usually fragrant, in axil of small (to rarely large and foliaceous) bract, with 2 (rarely several), bracteoles at or near base to near midpoint of pedicel. Calyx of 5 imbricate lobes, articulate with pedicel, persistent in fruit; corolla cylindrical to urceolate, with 5 short, imbricate lobes, (greenish) white to red, glabrous to densely covered with unicellular hairs abaxially, glabrous adaxially; stamens 10, in 2 whorls, inserted at base of corolla, the filaments flattened, geniculate, with long unicellular hairs, somewhat expanded near base, lacking spurlike appendages, the anthers more or less ovoid, minutely papillose to nearly smooth, lacking apical awns, dehiscing by large, introrse-terminal, elliptic pores, with white line or triangular patch of disintegration tissue on back of each lobe near apex, the pollen tricolporate, in tetrahedral tetrads, without viscin strands; stigma truncate to capitate, obscurely 5-lobed, minutely papillose, the style columnar, slightly swollen [not swollen] near apex, straight, with 5-fluted cen- tral canal, slightly to strongly impressed into apex of ovary and usually slightly exserted, the ovary superior, 5-locular, glabrous to densely covered with uni- cellular hairs (very rarely with few gland-headed hairs), with the placentae axile, subapical to central [basal] on persistent columella, slightly bilobed, bearing numerous anatropous ovules; nectariferous disc an enlargement of base of ovary wall, variously developed and lobed. Capsules loculicidal, subglobose, short-ovoid to ovoid, sutures unthickened (or very slightly thickened near apex), but not separating from valves at dehiscence (margins of valves whitish), placentae persistent on columella. Seeds very small, brown, scobiform, the testa thin, composed of single layer of very much elongated, thin-walled cells; embryo small, straight, more or less allantoid, white, with 2 small cotyledons, central in cross section, surrounded by fleshy endosperm. Germination epi- geal(?). 2n = 24 (Agarista populifolia). The name Agarista is taken from Greek mythology—the beautiful daughter of Clisthenes—in reference to the beauty of the flowers (see G. Don, 1834). DistriBuTION. South America—southeastern Brazil (from Bahia and Goias south to Rio Grande do Sul), Uruguay, northeastern Argentina, and Paraguay, along Andes from Bolivia and northern Peru to Colombia, mountains of southern Venezuela and adjacent Brazil (Para); Central and North America—- mountains of El Salvador and Honduras north to Mexico (Hidalgo, México, Jalisco), and on United States Atlantic Coastal Plain (from Florida to South Carolina); Map |. Also in central Africa, Madagascar, Réunion, and Mauritius (Sleumer, 1938, fig. 1). NuMBER OF SPECIES (TAXA): 30 (55). MEASUREMENTS AND TERMINOLOGY All measurements (except plant height, which was taken from information included on specimen labels or recorded in the field) included in the descriptions of species have been taken directly from dried herbarium material. The width 276 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 of the calyx lobes was measured midway between the apex and the point where they join the adjacent lobes; the width of all other structures was measured at the widest point. The values for inflorescence length refer to the range of variation in the longest inflorescence per specimen (and were determined by surveying total available herbarium material). As used here, “glabrous” and “‘pubescent” refer to the presence or absence of unicellular hairs; the presence, distribution, and density of multicellular, gland-headed hairs are described separately. SPECIMENS EXAMINED In the citation of specimens, abbreviations of institutions follow the seventh edition of Index Herbariorum (Holmgren, Keuken, & Schofield, 1981). In order to conserve space, only selected specimens have been cited. The locations of types cited but not seen have been taken from Sleumer (1959). l. I Key TO THE SECTIONS AND SPECIES OF AGARISTA Leaves with abaxial epidermis papillose; capsules with placentae basal; [Africa]; (sect. AGAU [30 Biel paierseenentiae Sah oes dance taenctenataraeyola tareeate Suet sachet ated aieke ota ee Ohi tee ame ey are a .] A. salicifolia (for key to infraspecific taxa, see Sleumer, 1938). Leaves with abaxial epidermis nonpapillose; capsules with placentae subapical to central; [Americas]; (sect. AGARISTA). 2. Leaves moderately to strongly adaxially folded. 3. Leaves 0.6-2.5(- as cm wide, narrowly to widely ovate, petiole 10-32 mm long. ...........................005. 27. A. glaberrima. 3. Leaves 0.1-0.5 cm wide, + linear, petiole 2-13 mm 4. Stems ewe inflorescences axillary or terminal racemes or panicles to 3-6 cm long, axis glabrous; leaves (1. we 7 cm long. Se ibe Bek Wek Sede boats thse mane Bere a ng Sil a Sos ea adel! wee nes Weare we ee Sed Be ES Soak ew sale ee tla ee 28. A. angustissima. 4. Stems glabrous to oe pubescent; inflorescences axillary racemes to 0.3-1.5 cm long, axis very slightly to see pubescent, AVES 152-3. 9( A OMLONG, 45 24 ce webww dd Pewee ulate Seas Sek eee eee es ce Pada eR E NG ead eee eereR ee eee 29. A. duartei. 2. Leaves plane to strongly on ie oe in bud. 5. Petiole slender, flexuous, elongate, 6-40 mm long. 6. Inflorescence axis glabrous; sane 4— 3 by 6-8 mm; leaves often slightly adaxially folded. ................. 27. A. glaberrima. 6. Inflorescence axis moderately to densely pubescent; a 3-5 by 4-6.5 mm; leaves + flat. 7. Leaves ovate; inflorescence axis with whitish hairs. ..........0 200000000 c ccc eee 25. A. boliviensis. aves oblong to ovate; inflorescence axis with cee Was: a cdaet oo oe eea see tt bas ae en tee 26. A. eucalyptoides. 5. Petiole stout and/or short, not flexuous, 0.5-15(—18) mm long. 8. Leaves 0.4-—2.5(-3.5) by 0.1-1.6(-2) cm Leaf margin + plane (to — lightly revolute, especially near base), lamina + flat. 10. Inflorescence axis glabro 11. Leaf blade with jength/width quotient > 1.8; corolla 7.5-13 mm long; filaments S-6.5 mm long. ... 16. A. pulchella. 11. Leaf blade with length/width quotient < 1.8; corolla 6-8.5 mm long; filaments 3.5-5 m 12. ee gland-headed hairs present on leaves and twigs; branches rigidly ce and wandlike, with modes usually to only ca. 0.5 cm long; leaves 0.4-1.3 cm wide. ..............0 0.0000 2000 20 5- 18. A. virgata. 12. Wee gland-headed hairs lacking; plants + widely to erectly branched, with internodes usually Ye 0.5-2 cm lene eaves (sh) COU WINGS 2.24 cadets a thuwaudekiedmectem sees as ietaeceetnedess 10. A. pulchra. 10. Inflorescence axis sparsely to densely hae axial leaf surface + densely pubescent. .......0..000.00 00 cc eee 11. A. subrotunda. 13. Atari leaf surface + a (but ites with few unicellular hairs on midvein). 14. Sepals 2.5-5 mm 14. Sepals 0.8-2.7 mm ie bee eee peed Ede Rese eisa ashes de Loe eae rk Te as ae ee 17. A. nummutaria. VLIslrdvov ‘dant [p861 LLC, 15. Capsules with placentae + centra 16. Inflorescence axis to 0.51. aes 5) cm long; leaf blade with length/width quotient usually > 3, cuneate to aa Catetal DASE. 3, straight sided, base cuneate to truncate, apex rounded- to bluntly acute-mucronate (never narrowly aaa PeHOle G6 MAN IONE, (5 24 aya 8 ose ee oa yen eda adwwy wey Bae oy Colee Cissihace Adin Se decd . A, minensis. 39. Inflorescence axis to 2-12 cm long; leaves various, but without above combination of characters. Leaves + ovate, 1.2—4.5(-5) cm long, margin usually convex-curved when viewed from above, base + cordate, petiole 1.5-5 mm long. 41. Lamina with abaxial glands lacking or very inconspicuous along midvein near base, ree coriaceous 16. A. pulc and flexible when dry; [Parana, Santa Catarina, Sao Paull). esis aetnte date cee hella. 41. Lamina g g near base, thickly coriaceous and quite inflexible when dry; [Minas Gerais]. 42. Inflorescence axis + ecu pubescent. .......0 000 eee 9. A. coriifolia. 42. INMOPESCENCS ANAS PIADLOUSS 5 xcd-csin. be Me pew ck candle neh wads bok eee eeadtc 10. A. pulchra. 40. Leaves ovate to elliptic or oblong, a 12.5 cm long, margin often + straight when viewed from above, base cuneate to slightly cordate, petiole 2 TS MN IONGS opin kind nc ehe beeen eareka's 14. A. oleifolia. 37. Capsules with placentae + subapical; seeds 1-3.2 mm lon Inflorescence axis usually moderately to densely covered with ferrugineous unicellular hairs. ................. 43. ig as axis glabrous to densely pubescent, hairs not ferrugineous. rescence axis to 0.5—3 cm lon 45. pre uniformly ovate, apex a acumin Leaf margin conspicuously aes eee Wee s Goce ta fae ee added 25. A. boliviensis. 46. Leaf entire to serrate, margin not aie ab crisped/undulate. WOLAYOUUV GTIONYV AHL AO TVNUNOL ¢9 “10A] as ~ Cc) i) ko} wn = oO wm ie — ie by 3 =| a i=) ge =. = =e toma =. O > o =] o a I+ = rat) al Mt chad im] [o) & i’) =) oO jon = = a 7) m ive] ol ro) a= aN ~ . Capsules 3-4.5 mm long, with thin, + smooth walls; [North and Central Ameri | . Ovary very sparsely to densely pubescent; pith nonseptate to septate; bark sanenily and deeply Miro wed {COPY 5j:0- 8 es oe, eed oh as ene Abigale baie arid eRe Gna SS 6. A. mexicana. 48. Ovary glabrous; pith always clearly septate; bark variable. 49. Calyx glabrous on abaxial surface; pedicels glabrous; corolla 8.5-13 mm long; leaves entire; bark furrowed, corky; [Mexico]. .................000000005- 7. A. sleumeri. 49. Calyx sparsely to moderately pubescent on abaxial surface; pedicels sparsely to mod- ria! pubescent; corolla 6.5-9.5 mm long; leaves entire or serrate; bark very shallowly furrowed, not corky; [SE United States]. .....................005. 8. A. populifolia. 45. rae of oe olan aes from ovate to elliptic or oblong (rarely obovate), apex acute- or rounded- mucronate to a 50. Che iene (ees leaves with apex acute- or rounded-mucronate to acuminate, margin convex-curved to + straight when viewed from above; [SE Brazil]. 51. Leaves ovate to elliptic Hen oblong), margin very slightly revolute from base to near apex; Giianabara. Rio: de Janette]: vi csce era even winenenr ee cece eines see si ines 24. A. uleana. 51. Leaves ovate to alicueieblote ees at least few per plant + parallel sided, wae plane to very slightly revolute at base only; [Rio Grande do Sul to Parana]. ....... 23. A. niederleinii. Capsules + smooth; leaves with apex elongate-mucronate to short-acuminate, margin convex-curved n viewed from above; [chiefly N South America]. ..................000000005- 4. A. duckei. 44. je ee axis to 3-15 cm 52 orescence axis with hairs to 0.15-0.4 mm long, straight to curved; calyx lobes 0.7-1.8 mm long; seal (3.7-)4-9 mm long; leaves thinly coriaceous, flexible when dry; [N Andes]. _1. A. albiflora. 52. Inflorescence axis glabrous or with hairs to 0.05-0.15 mm long, curved; calyx lobes 120 7 mm long; corolla 6.5-11 mm long; leaves thickly coriaceous, rather inflexible when dry; [SE Brazil]. ........... i fa dreds 2h ais thes & Apes ge Geek ete BE at EN RIL Ea a as ee a ee et pears ee es ._ A. coriifolia. au, oO VLsIavov ‘dant [p861 I8¢ 282 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Agarista D. Don ex G. Don sect. Agarista Leucothoé D. Don sect. Agastia DC. Prodr. 7: 603. 1839. Leucothoé D. Don subg. Agarista (D. Don ex G. Don) Drude in Engler & Prantl, Nat. Pflanzenfam. 4(1): 42. 1889. LecrorTyPE SPECIES: oe nummularia (Cham. & Schldl.) G. Don. Agarista D. Don ex G. Don sect. Leucothoides Niedenzu, Bot. Jahrb. 11: 186. 1890. LECTOTYPE SPECIES: Agarista ee (Cham.) G. Don (= Agarista chlorantha (Cham.) G. Don Agarista D. Don ex G. Don sect. Euagarista Niedenzu, Bot. Jahrb. 11: 186. 1890, nomen illegit. Twigs gas nonchambered to clearly chambered, Calluna- ae pith. Leaves with abax id ill ada y with divided cells (see Stevens, 1970). Style usually swollen toward apex. “Capsules with placentae subapical to + central; seeds 0.5—3(-3.5) mm long. NUMBER OF SPECIES (TAXA): 29 (35). 1. Agarista albiflora (B. Fedtsch. & Basilevsk.) Judd, comb. nov. FIGURE 2, d. Leucothoé Ke B. Fedtsch. & Basilevsk. Not. Syst. Herb. Hort. U.S.S.R. 6: eee E: Colombia, eae near Soata, April 1843, Linden 300 (holotype, LE; otypes, BR, F!, G, K!, NyY!, p!, us!, w). ieee ies mee Sane Notizbl. Bot. Gart. Berlin 12: 479. 1935. Type: Co- lombia, Magdalena, Sierra Nevada de Santa Marta, ‘El Manon,” ca. 2100 m alt., 26 Feb. 1928, Schultze 1113 Pes B (destroyed)). Leucothoé andina Sleumer, ibid. 478. Tyre: Peru, Amazona s, E of Chachapoyas, 2000 m alt., 19 July 1904, Weberbauer 4362 ene B TV Gestoicd: isotypes, ny(fragment)!, usm). Shrub to 3 m tall. Twigs with or without scattered gland-headed hairs, oth- erwise moderately pubescent, with + nonchambered pith. Buds to ca. 1.5 mm long. Leaves alternate; petiole 2-8 mm long; blade revolute in bud, ovate to elliptic, 1.6—7 by 0.8-2.7 cm, flat to slightly (strongly) abaxially curved, cori- aceous, the apex acute- to rounded-mucronate, the base widely cuneate to cordate, the margin entire (slightly undulate), plane to very slightly ee revolute, the adaxial surface with or without gland-headed hairs on midvein, otherwise sparsely to moderately pubescent on midvein (also sparsely ne on lamina and along margin), the abaxial surface with or without gland-headed hairs on midvein, otherwise very sparsely to moderately pubescent on midvein, usually with few to many inconspicuous glandular dots along midvein. Inflo- rescences axillary racemes to 3—10 cm long, the axis with or without scattered gland-headed hairs, otherwise sparsely to moderately pubescent. Pedicels 2— 7.5 mm long, with or without gland-headed hairs, otherwise moderately pu- bescent; bracteoles 2, opposite to subopposite, basal or nearly so, narrowly triangular, to ca. 0.8 mm long; bracts to 1 mm long. Calyx lobes triangular with acuminate apices, 0.7-1.8 by 0.6-1.1 mm, abaxial surface sparsely to moderately pubescent; corolla cylindrical to urceolate-cylindrical, 4-9 by 2- 3.5 mm, white (red tinged at tip, or red), abaxially glabrous; filaments 2.5-5 mm long, anthers |-1.5 mm long; ovary sparsely to densely pubescent. Capsules 1984] JUDD, AGARISTA 283 | Figure 2. a, b, Agarista mexicana var. pinetorum: a, Williams & Molina 14000; b, Villarreal de Puga 4761, abaxial leaf surface light due to dense layer of unicellular hairs. c, A. revoluta (Rose & Russell 20687). d, A. albiflora (Jorgensen & Prieto JP-55). Scale = 2 cm. 284 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 T T 80 70 60 50 Map Distribution of Agarista albiflora (dots), A. subcordata (triangle), A. braca- morensis (square), and A. duckei (circles). subglobose to short-ovoid, 3—5.5 by 4-7.5 mm, placentae subapical; seeds | .5— 2 mm lon DiIsTRIBUTION AND ECOLOGY. Northern Andes from Venezuela and Colombia south to Peru (Map 2). Moist montane forests, dwarf forests, rocky areas with scattered shrubs; 1800-2500 m alt. Flowering April to September. REPRESENTATIVE SPECIMENS. Colombia. Boy Aca: Villa de Leiva, La Candelaria, Cleef 359 (L). MAGDALENA: San Sebastian de Rabago, Romero-Castafeda 867 (co). Venezuela. Meripa: San Antonio, Jahn 1086 (us); Pueblo Nuevo, Lépez-Palacios 768 (Ny). TACHI- RA: above La Grita, slopes below Paramo de la Negra, Stevermark 57108 (A, F). Ecuador. Loysa: Hacienda Guaycopamba on Rio Guaycopamba, 22 km S of Vilcabamba, Fosberg & Giler 23072 (COL, NY, US); W slope of Cordillera Condor, Huilcabamba, NW of Nudo de Sabinilla, 54 km S of Loja, Jorgensen & Prieto Jp-55 (FLAS, NY, 8). Peru. CAJAMARCA: Jaén, El Parco, Friedberg 665 (L). Agarista albiflora, a taxon that is rather variable in leaf size and shape, and in the degree to which the margins are revolute, is probably most closely related to A. subcordata and A. bracamorensis. It can be distinguished from the former 1984] JUDD, AGARISTA 285 by its often larger leaves and its inflorescence axes with usually elongated, straight to curled, unicellular hairs, and from the latter by its much larger, usually nonrevolute leaves and its often longer pedicels. Agarista albiflora may also be confused with A. revoluta and A. duckei, or A. mexicana, A. sleumeri, and A. populifolia (see key for distinguishing characters). Agarista albiflora and A. subcordata may be slightly sympatric since both have been collected in the Chachapoyas region; the mechanisms isolating these two very similar species are in need of field study. 2. Agarista subcordata (Dunal) Judd, comb. nov. Gaylussacia subcordata Dunal in DC. Prodr. 7: 557. 1839. Leucothoé subcordata Dunal) Sleumer, Bot. Jahrb. 78: 461. 1959. Type: Peru, Amazonas, Chachapoyas, Matthews s.n. (holotype, G; fragment and photo of holotype, F!; isotypes, E!, K!, s!). Shrub. Twigs sparsely to very sparsely pubescent, with + nonchambered pith. Buds to ca. 1.8 mm long. Leaves alternate; petiole 1-2.5 mm long; blade revolute in bud, ovate to elliptic, 1-3.1 by 0.6-1.3 cm, flat to slightly abaxially curved, coriaceous, the apex acute- to rounded-mucronate, the base rounded to slightly cordate, the margin entire (slightly undulate), the adaxial surface glabrous to sparsely pubescent on midvein, the abaxial surface glabrous to sparsely pubescent on midvein, often with few to several inconspicuous glan- dular dots along midvein. Inflorescences axillary racemes to 5—10 cm long, axis sparsely to moderately pubescent. Pedicels 3.5—7 mm long, sparsely to mod- erately pubescent; bracteoles 2, at or near base to within lower 3 of pedicel, narrowly triangular, to ca. 1.1 mm long; bracts to 1.5 mm long. Calyx lobes triangular with acuminate apices, |-1.5 by 0.9-1.5 mm, the abaxial surface glabrous to very sparsely pubescent, especially near base; corolla cylindrical to urceolate-cylindrical, 6-7.5 by 3-3.5 mm, white, abaxially glabrous; filaments ca. 3.5 mm long, anthers ca. 1.3 mm long; ovary glabrous to very sparsely pubescent near apex. Capsules subglobose to short-ovoid, 4-5 by 5.5-6.5 mm, placentae subapical; seeds not seen. DisTrRiBUTION. Andes, endemic to Chachapoyas region of northern Peru, ca. 2700 m alt. (Map 2). REPRESENTATIVE SPECIMENS. Peru. AMAZONAS: Chachapoyas, Guancas, Matthews 1635 (E, GH, K, Pp); Chachapoyas, Williams 7549 (F). The poorly known Agarista subcordata is probably closely related to both A, albiflora and A. bracamorensis. All three taxa are limited to the northern Andes, but A. albiflora is by far the most widely distributed. Agarista subcordata can be readily distinguished from 4. albiflora by its often smaller leaves and its inflorescence axis with short, curled hairs, and from 4. bracamorensis by its wider, less strongly revolute leaves and its longer pedicels. 3. Agarista bracamorensis (Humb., Bonpl., & Kunth) G. Don, Gen. Syst. 3: 837. 1834. Andromeda bracamorensis Humb., Bonpl., & Kunth, Nov. Gen. Sp. Pl. 3: 225. ¢. 263. 1818. Leucothoé bracamorensis (Humb., Bonpl., & Kunth) DC. Prodr. 7: 603. 1839. 286 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Type: Peru, Cajamarca, prov. Bracamora, near S. Felipe and Paramo de Yamoca, ca. 2350 m, Bonpland 3564 (holotype, p!; fragment and photo of holotype, F!). Shrub or small tree. Twigs sparsely pubescent, with obscurely chambered pith. Buds to ca. 0.5 mm long. Leaves alternate; petiole 1.5-4 mm long; blade revolute in bud, ovate, 0.8-1.5 by 0.25-0.7 cm, moderately abaxially curved, coriaceous, the apex acute-mucronate, the base rounded, the margin entire, strongly revolute, the adaxial surface sparsely pubescent on midvein, the abaxial surface essentially glabrous, often with few inconspicuous glandular dots along midvein. Inflorescences axillary racemes to ca. 5 cm long, axis sparsely to moderately pubescent. Pedicels |1.5-3 mm long, sparsely to moderately pu- bescent; bracteoles 2, opposite, nearly basal, narrowly triangular, to ca. 0.6 mm long; bracts to | mm long. Calyx lobes triangular with acute to acuminate apices, 0.5-1.5 by 0.4-0.6 mm, abaxial surface glabrous; corolla + cylindrical, 4.5-6 by 2-3.5 mm, white, abaxially glabrous; filaments ca. 3.5 mm long, anthers ca. 1.3 mm long; ovary glabrous. Capsules short-ovoid, 4-4.5 by 4.5- 5 mm, placentae subapical; seeds ca. 1.5—2 mm lon DistRIBUTION. Andes, northern Peru, Bracamora region, ca. 2350 malt. (Map 2). REPRESENTATIVE SPECIMENS. Known only from type collection. This rare and little-known species is probably closest to Agarista albiflora and 4. subcordata, two other Andean species. It is readily separated from these allies by its very small, strongly revolute leaves. It is also superficially similar to the revolute-leaved Brazilian species 4. organensis, A. chlorantha, A. his- pidula, and A. ericoides but is quickly distinguished from all of them by its combination of round-based leaves, short pedicels, and short calyx lobes. 4. Agarista duckei (Huber) Judd, comb. nov. se rece Huber, Bull. Soc. a Genéve, II. 1: 246. fig. J. 1909. Type: Brazil, A, mpis arenosis ad ripam orient. fl. Yamunda, pr. Faro, 27 Aug. 1907, Ducke 25.26 holotype, MG; tae and photo of holotype, F!; isotype, G-Boiss.). Leucothoé venezuelensis A. C. Smith, Contr. U. S. Natl. Herb. 29: 335. 1950. Type: Venezuela, Bolivar, Gran Sabana, Cerro Akurima, 20 March 1946, Tamayo 3236 (holotype, us!; photo of holotype, Ny!; isotype, F!). —_ Shrub to tree to 6 m tall, with roughly furrowed bark. Twigs sparsely to densely pubescent, with nonchambered pith. Buds to ca. 1.5 mm long. Leaves alternate; petiole 2-6(—7) mm long; blade revolute in bud, elliptic to slightly ovate (obovate), (1.5-)2-5.7 by 0.5-2.1 cm, nearly flat to strongly abaxially curved, coriaceous, the apex elongate-mucronate to short-acuminate, the base narrowly to widely cuneate or rounded, the margin entire, plane to strongly revolute, the adaxial surface sparsely pubescent on midvein, especially prox- imal portion (with few scattered hairs on proximal portion of lamina), the abaxial surface sparsely to moderately pubescent on midvein, with few to several very inconspicuous glandular dots along midvein. Inflorescences axil- lary racemes to 0.5—1.5 cm long, axis densely pubescent. Pedicels 2.5—12 mm 1984] JUDD, AGARISTA 287 long, moderately to densely pubescent; bracteoles 2, alternate, from nearly basal to within lower '4 of pedicel, triangular to narrowly so, to ca. 1.2 mm long; bracts to 1.2 mm long. Calyx lobes triangular with acuminate apices, 0.9-2.3 by 0.7-1 mm, abaxial surface moderately to densely pubescent; corolla urceo- late to cylindrical, 6-8 by 3-4.5 mm, white (rarely pink), abaxially glabrous; filaments 3.5-4.7 mm long, anthers 0.9-1.1 mm long; ovary sometimes with scattered gland-headed hairs, otherwise sparsely to densely pubescent. Capsules ovoid to globose or subglobose, 5-6 by 6-8 mm, valves rarely white bordered, placentae subapical; seeds 1.5-3 mm long. DISTRIBUTION AND ECOLOGY. Southern Venezuela and western Para and Mato Grosso, Brazil (Map 2). Thickets and forests on sand, open sandy areas, bush forest and savannas on white sand, river and lake margins, rocky slopes and plateaus; ca. 100-1400 m alt. Flowering May through August (October and December) : SPECIMENS. Brazil. MATO Grosso: Sararé, ance eti Pires & Santos 16397 ice: ao ParA: E of Faro, Ducke 5708 (Ny, P, 8, US; photos, F, re Monte Alegre, Alto de Serra do Ereré, Lima 53-1609 (k). Venezuela. rene : Sabanito orocoto, right bank of Rio Orinoco, 2 km below mouth of Rio Aiabape: i ‘urdack & Adderley 42692 (F, GH, NY, UC, US, VEN). BoLivar: Gran Sabana, Canaima Lagoon, Avensa Camp, Ehrendorfer 74103-1b (ven), Canaima, Lopez-Palacios 3029 (L, NY), Canaima, between hotel and airport, Stevermark 106370 (F, MO, NY, VEN); Carretera El Dorado to Santa Elena de Uairén, km 198 S of El Dorado, plateau above Lamé-mert, Stevermark et al. 106642 (vEN); top of rocky slope, Steyermark et al. 106634 (Ny). The closest relative of Agarista duckei seems to be the geographically sep- arated A. revoluta (see MAP 5). The strong similarities between the two species were also noted by Sleumer (1959). In both species the leaf blade is often strongly to moderately abaxially curved. Agarista duckeiis readily distinguished from A. revoluta by its shorter inflorescences, its short-acuminate to elongate- mucronate leaf apices, and its usually white flowers. Agarista duckei is also related to (and may sometimes be confused with) the Andean species 4. al- biflora, see key for distinguishing characters. Albert C. Smith (1950) distinguished Leucothoé venezuelensis (populations in Venezuela) from L. duckei (populations in state of Para, Brazil) on the bases of petiole length, leaf shape and base, calyx-lobe apex, and corolla shape. However, the variation in these characters is so slight that separating the plants into two species is quite arbitrary. Thus, the two are considered to be conspe- cific. The large disjunction separating the Mato Grosso population from similar plants in southern Venezuela is puzzling. Additional localities probably exist; more collecting is evidently needed. The pollen morphology of this species (Venezuelan populations) has been studied by Maguire, Steyermark, and Luteyn (1978). 5. Agarista revoluta (Sprengel) J. D. Hooker ex Niedenzu, Bot. Jahrb. 11: 236. 1889 Shrub or small and often gnarled tree to 3(-6) m tall, with roughly furrowed bark. Twigs sparsely to densely pubescent, with nonchambered pith. Buds to 288 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 ca. 1.1 mm long. Leaves alternate; petiole 2-7 mm long; blade revolute in bud, ovate to elliptic, 1.3-4.5 by 0.5-2.2 cm, slightly to strongly abaxially curved (rarely nearly flat), coriaceous, the apex obtuse- to retuse-mucronate (rarely acute-mucronate), the base rounded (to very slightly cordate), the margin entire, slightly to strongly revolute, the adaxial surface sparsely to moderately pubes- cent on proximal portion of midvein (with few hairs on adjacent portion of lamina); abaxial surface sparsely pubescent on midvein or only on proximal portion, or densely pubescent on lamina and midvein, with few inconspicuous glandular dots along midvein. Inflorescences axillary racemes to (1-)1.5-6.5 cm long, axis moderately to densely pubescent. Pedicels 3-1 1 mm long, sparsely to densely pubescent; bracteoles 2, opposite to alternate, from basal to near midpoint of pedicel, narrowly triangular, to ca. | mm long; bracts to 1.3 mm long. Calyx lobes triangular with acuminate apices, 0.9-1.6 by 0.6-1.5 mm, abaxial surface sparsely to densely pubescent; corolla + cylindrical, 6-9 by 3.5-5 mm, red to white, abaxially glabrous or with very few unicellular hairs near apex to moderately pubescent; filaments 4.5-5.5 mm long, anthers 1-1.2 mm long; ovary very sparsely to densely pubescent. Capsules ovoid to short- ovoid or subglobose, 4-5.5 by 5-7 mm, valves often slightly white bordered, placentae subapical; seeds 2-3.5 mm long. DiIsTRIBUTION. Southeastern Brazil, along coast from Bahia south to Estado do Rio. KEY TO THE VARIETIES OF AGARISTA REVOLUTA 1. Abaxial surface of lamina glabrous, but sparsely pubescent on midvein, especially proximal portion; corolla glabrous or with very few unicellular hairs near apex Sopa tes Raya eoseteas teeta one a sapere ste ye yehes oe ede raat ad to eee seen hae dk os Ede Sa. var. revoluta. |. Abaxial surface of lamina d yp t; corolla moderately pubescent, especially NGAI ADCXy, sei Sie eae aie sath te is earaearad whan Ga eeded Oa knees Miawies 5b. var. velutina. 5a. Agarista revoluta (Sprengel) J. D. Hooker ex Niedenzu var. revoluta IGURE 2, C. Andromeda revoluta Sprengel, Neue Entdeck. 2: 131. 1821. Agarista sprengelii G. Don, Gen. Syst. 3: 837. 1834, nomen superfl. Lewcothoé revoluta (Sprengel) DC. Prodr. 7: 604. 1839. mera revoluta (Sprengel) DC. var. se/lowii Meissner in Martius, Fl. Brasil. 7: 160. 1863, nomen superfl. Tyre: Brazil, Estado do Rio, Restinga de Cabo Frio, Se/low s.n. a B teenie isotype at B! selected as see remaining isotypes, BM!, BR, E!, , w; photos of isotype, F!, sndomea bales a Nees, Flora 4: 297. 1821. Type: Brazil, Estado do Rio, Cabo ied 13, 14 (not seen, probably at BR, GOET; see Sleumer, 1959). i ea cee DC. var. salzmannii DC. Prodr. 7: 604. 1839 (= L. bahiensis var. bahiensis; chosen as nominate variety by Sleumer, 1959). Leucothoé revoluta (Spren- gel) DC. var. salzmannii (DC.) Meissner in Martius, Fl. Brasil. 7: 160. 1863. Type: Brazil, Bahia, 1830, Salzmann 314 (lectotype, G; isolectotypes, Nv(fragment)!, p!: unnumbered Salzmann collections, possible isolectotypes, G, GH!, K!, Mo!; photos of unnumbered eee ae ion, F!, G, GH! sr ete bahie DC. v r, blanchetii DC. Prodr. 7: 604. 1839. Leucothoé revoluta (Spre vel) DC. ¥ var. Fuache (DC.) Meissner in cours aes Type: Brazil, Bahia eae cannes Blanchet 1680 (holotype, G; isotypes, KI, WwW). 1984] JUDD, AGARISTA 289 Andromeda nitida Vell. Conc. Fl. Flum. 174, 1825, a later homonym of A. nitida Bartram ex Marsh. Arbust Am. 8. 1785 = Lyonia lucida (Lam.) K. Koch. Leucothoé bahiensis DC. var. arrabidae DC. Prodr. 7: 604. 1839. Leucothoé revoluta (Sprengel) DC. var. arrabidae (DC.) Meissner in Martius, Fl. Brasil. 7: 160. 1863. Type: lec- totype here designated as illustration in Vell. Conc. Fl. Flum. ic. 4: ¢. 94. 1835 Twigs sparsely to densely pubescent. Leaves with adaxial surface sparsely pubescent on proximal portion of midvein; abaxial surface sparsely pubescent on midvein, especially proximal portion, otherwise glabrous. Pedicels 3-8.5 mm long, sparsely to densely pubescent. Calyx lobes with abaxial surface sparse- ly to densely pubescent. Corolla abaxially glabrous or with very few unicellular hairs near apex. Ovary very sparsely to sparsely pubescent. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, along coast from southern Bahia south to Estado do Rio (Map 5). Coastal scrub on wet to dry sand (sometimes intermixed with sedge meadows), sand dunes; near sea level (rarely inland to 1000 m alt.). Flowering July through October. REPRESENTATIVE SPECIMENS. Brazil. BAHIA: Santa Cruz Cabralia, Belém & Pinheiro 2544 (c, L, NY); Pérto Seguro, Duarte 6144 (L, M, Ny, Us); between Alcobaga and Caravelas on BA 001, 20 km S of Alcobaca, Harley 18047 (Ny); 7 km NW of Mucuri, Mori et al., Herb. Cent. Pesquisas do Cacau 10474 (Ny). Estapo bo Rio: Itapua, Athayde III- 1961 (L); Macahé, Restinga de Cabiuna, Brade 15 78 I (3); Cabo Frio, Duarte & Pereira 5 793 Segadas-Vianna et al. I-949 (1); Restinga de Itaipu, Sucre 76/1 (Ny); Restinga de Cabo Frio, Ule 4752 (HBG). Minas Gerats: Serra Sapucai, municipio de Jequitinhonha, Ma- galhaes 17459 (L). 5b. Agarista revoluta (Sprengel) J. D. Hooker ex Niedenzu var. velutina Judd, var. nov Varietas haec ab Agarista revoluta var. revoluta differt in foliis cum pagina abaxiali dense velutina, pedicellis longioribus, corollis et ovariis pubescentibus. Twigs densely pubescent. Leaves with adaxial surface sparsely to moderately pubescent on midvein and adjacent lamina; abaxial surface velutinous on lam- ina and midvein. Pedicels 6-11 mm long, densely pubescent. Calyx lobes with abaxial surface densely pubescent. Corolla abaxially moderately pubescent, especially near apex. Ovary + densely pubescent. Type. Brazil, Bahia, municipio do Salvador, Dunas do Abaeté, 29 Aug. 1976, J. Silva Aratjo et al. 49 (holotype, Ny!; isotype, CEPEC). DISTRIBUTION AND ECOLOGY. Eastern Brazil, on coast near Salvador, Bahia (Map 5). Coastal scrub. Flowering August and September. REPRESENTATIVE SPECIMENS. Known only from type collection. Agarista revoluta, a distinct species with variably revolute, moderate-sized, obtuse- to retuse-mucronate leaves, is probably allied to the geographically 290 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 65 separated A. duckei (see Map 2). It is readily distinguished by its longer inflo- rescences and its more abruptly and shortly te leaves. Some individuals of A. albiflora, an Andean species, may be confused with A. revoluta, however, the former taxon has leaves that are usually less abruptly mucronate and only infrequently revolute, inflorescence hairs that are longer, and capsule valves that are never bordered. Agarista revoluta is divided here into two geographically isolated varieties (Map 5) separable primarily by the indumentum on the abaxial leaf surface. Variety ve/utina, discovered only recently, may be more widespread, and fur- ther collecting is needed along the coast of Bahia. This taxon is unique in that it is the only species of Agarista known to occur in coastal “restingas.”’ 6. Agarista mexicana (Hemsley) Judd, Jour. Arnold Arb. 60: 495. 1979. Shrub or tree to 8(-11) m tall, with thick, corky, deeply furrowed bark. Twigs very sparsely to densely pubescent, with nonchambered to clearly chambered pith. Buds to ca. 1.5 mm long. Leaves alternate (to subopposite or nearly 3-whorled at some nodes); petiole 2-12.5 mm long; blade revolute in bud, ovate, 2—7.5(-9) by 0.8-2.5(—3.2) cm, flat, coriaceous, the apex acuminate, the base narrowly cuneate to rounded, the margin entire (slightly undulate), plane to very slightly revolute near base, the adaxial surface glabrous to densely pubescent on midvein (with few unicellular hairs on proximal portion of lam- ina); abaxial surface essentially glabrous to densely pubescent on midvein, otherwise glabrous to densely pubescent, with few to several inconspicuous glandular dots along midvein. Inflorescences (fasciclelike) axillary racemes to 0.5—2.5(—4) cm long, axis sparsely to densely pubescent. Pedicels 2-8 mm long, very sparsely to densely pubescent; bracteoles 2, alternate to opposite, from nearly basal to within lower ' (rarely to midpoint) of pedicel, triangular to narrowly so, to ca. 0.9 mm long; bracts to 1 mm long. Calyx lobes triangular with acuminate apices, 0.9-1.8 by 0.5-2 mm, abaxial surface very sparsely to densely pubescent; corolla cylindrical, 6-9.5 by 2.5—-5 mm, white, abaxially glabrous to sparsely pubescent (especially near apex); filaments 4.5-6 mm long, anthers |-1.8 mm long; ovary very sparsely to densely pubescent. Capsules subglobose to short-ovoid, 3-4.5(-5) by 5-7 mm, placentae subapical; seeds 1.5-2.7 mm long. DisTRIBUTION. Mountainous areas of Mexico and Central America, from Ve- racruz and Jalisco south to Honduras and El Salvador. COMMON NAMES. Cachimbo (Honduras, see Standley & Williams, 1952); que- manote, pellejo de lagarto, nacahuite (Mexico). KEY TO THE VARIETIES OF AGARISTA MEXICANA —_ . Abaxial surface of sua ae to occasionally sparsely pubescent, hairs not obscuring abaxial epidermis. .................00.......000.. a. var. mexicana. Abaxial surface of Cane eee to occasionally moderately pubescent, hairs + obscuring epidermis. ......0..00.00. 0000 c eee tees 6b. var. pinetorum. 1984] JUDD, AGARISTA 291 6a. Agarista mexicana (Hemsley) Judd var. mexicana Andromeda mexicana Hemsley, Biol. Centr.-Am. Bot. 2: 282. 1881. Leucothoé mex- icana (Hemsley) Small, N. Am. Fl. 29: 57. 1914. Type: Mexico, Oaxaca, Sierra San Pedro Nolasco, Jiirgensen 866 (holotype, kK; isotype, G!) Twigs very sparsely to densely pubescent. Leaf blades with abaxial surface glabrous to occasionally sparsely pubescent, epidermis not obscured. Ovary very sparsely to densely pubescent. DISTRIBUTION AND ECOLOGY. Mountainous areas of Mexico and Central Amer- ica, from states of México, Guerrero, Oaxaca, and Veracruz south to Honduras and El Salvador (Map 3). Dry ridges, Pinus and/or Quercus forests, Pinus, Quercus, and Liquidambar forests with Ulmus, Zinowiewia, Weinmannia, and Styrax, montane mixed forests, disturbed openings, and forest margins; fre- quently in sandy soil; 170-2500 m alt. Reported from forests of Pinus teocote, P. rudis, and P. oocarpa by Gémez-Pompa (1973). Flowering mid-February through April (early May). REPRESENTATIVE SPECIMENS. Mexico. CuiApas: SE of Cerro Baul on border with Oaxaca 16 km NW of Rizo de Oro, municipio de Cintalapa, Breedlove & Smith 31375 (MEXU, mo); SW of Tal-tenango, along rd. to Finca Pousia, Miranda 6945 (MEXU, US). GUERRERO: Agua de Obispo, near 17°20'N, 99°34’ W, Kruse 630 (ENCB, MEXU);, Taxco, Miranda 3074 (mEXU). México: dist. Temascaltepec, Guayabal, Hinton 3377 (Gu, NY, US); Nanchititla, Hinton 3415 (Gu, Ny, Us); Temascaltepec, Hinton 3528 (Gu, Ny, UC, US); dist. Sultepec, Almoloya, Hinton et al. 7450 (BM, F, G, GH, MO, NY, US); Cerro de Ma matla, Zacualpan, Tejupilco, Rzedowski 22122 (wis). Oaxaca: Zempoaltepetl, between Santa Maria and Mitla, Camp 2761 (GH, Ny, 8, UC); Chavela, — ae NY, US); ast. Choapam, Yaveo, trail to Arroyo Culebras, Mexia 9157 (Bb, F, G, MO, NY, S, UC, US); Teotalcingo to Choapan, Reko 4080 (us), Choapan to aes Reko 4116 (us); between Rio Grande and Niltepec, Yolocotzi X-1230 & Sharp 46232 (GH, MEXU). VERACRUZ: Cerro Monte de Oro, Dorantes et al. 856 (F, MEXU); Falda del Cerro Azul, a eles Limon-- Cerro Azul, Dorantes et al. 1114 (mexu); along trail to Santa Marta, 2 km N of San Fernando, municipio Soteapan, Nee et al. 24729 (F, FLAS), Ocozotepec, 6 km NW of Soteapan, Sousa 3253 (Fr, MEXU); | km SW of Ocozotepec, Sousa 3532 (F, MEXU). Belize: Hidden Valley Falls Rd., eal g.n., 26 Dec. 1973 (FLAS). Guatemala: Zamorora, dept. Santa Rosa, Heyde & Lux 4530 (F, G, GH, NY, Us); Sierra de las Minas, near San Geronimo, dept. Baja Verapaz, Kellerman 6636 ‘S us); above Morazan, El Progreso, Sharp 4633 (GH, MEXU); between Finca San José and Montana Nube, 1.5 mi SE of Concepcidn de las Minas, dept. Chiquimula, eee 30857 (F, NY); Montafia Castilla, vic. of Montafia Cebollas, 3 mi SE of Quezaltepeque, dept. Chiquimula, Steyermark 31303 (F); Sierra de las Minas, dept. Zacapa, Volcan de Monos, Steyermark 42317 (r, Ny); Sierra de las Minas, near Finca Piamonte, dept. El Progreso, Stevermark 43427 (F, ny); Aguacate, dept. Jalapa, Williams 13172 (F, Gu); near Soledad, dept. Jalapa, Williams 14239 (Fr). Honduras: dept. Comayagua, near Siguatepeque, Allen 6219 (F); dept. Intibuca, Cascada de eae Molina 6362 (F); dept. Yoro, above Los Flores, vic. 0 Coyoles, Yuncker et al. 8172 (BM, F, G, GH, NY, US). E] Salvador: dept. Chalatenango, La Reina, Calderon (F, us); dept. Santa Ana, Cerro Miramundo, NE of Metapan, Carlson 956 (F, Uc); dept. Chalatenango, S of La Palma, Lagos & Weberling 256 (L, M); dept. Morazan, easternmost peak, Montes de Cacaguatique, lat. 13°46'N, long. 88°13’W, Tuck- er 698 (G, NY, P, UC, US). 292 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 aT 1 A 1 4 Map 3. Distribution of Agarista mexicana var. mexicana (circles), A. mexicana var. pinetorum (dots), and A. sleumeri (squares). Localities with both A. mexicana var. mex- icana and intermediates between this taxon and var. pinetorum indicated by half-closed circles 6b. Agarista mexicana (Hemsley) Judd var. pinetorum (Standley & Williams) Judd, Jour. Arnold Arb. 60: 495. 1979. FIGURE 2, a, b. Leucothoé pinetorum Standley & Williams, Ceiba 3: 54. 1952. Leucothoé mexicana (Hemsley) Small var. pinetorum eae & Williams) Sleumer, Bot. Jahrb. 78 466. 1959. Type: Honduras, El P , Manzaragua, 1400 m alt., 4 April las Williams & Molina ee Gee. Te photos of holotype, F!, Gu!, Ny!, uc! isotypes, GH!, MEXU!, Mo!, Us!). Twigs moderately to densely pubescent. Leaf blades with abaxial surface densely (to moderately) pubescent, epidermis obscured or nearly so. Ovary moderately to densely pubescent. DISTRIBUTION AND ECOLOGY. Mountainous areas of Mexico (states of Jalisco, Guerrero, and Chiapas), Belize, and Honduras (Map 3). Rocky, brushy hillsides, Pinus and/or Quercus forests, Pinus savannas, Pinus, Quercus, and Liquid- ambar forests, and montane mixed forests; often in sandy soil; 500-2000 m alt. Flowering March through April (early May). REPRESENTATIVE SPECIMENS. Mexico. CHIAPAS: E base of Cerro Tres Picos near Cer: Bola, logging rd. SW of Colonia Agronomos Méxicanos, municipio Villa Corzo, pe . Thorne 30037 (DUKE, ENCB, F, MEXU, MO). GUERRERO: Mina, Manchon dist., anon 0082 (c, F, G, GH, M, MO, NY, UC); Armenia-Zoyate, Mina, Hinton 10159 (c, ENCB 1984] JUDD, AGARISTA 293 G, GH, M, MO, NY, UC, US); Chilacayote-La Soledad, Mina, Hinton et al. 14199 (F, GH, MO, NY, UC, us); Petlacala, Mina, Hinton 14205 (Ff, Mo, NY, US). JALISco: La Venta. municipio de Zapopan, Didz-Luna 87 ee B); Venta del Astillero, La Primavera, Vi/la- rreal i Puga 150 (ence); Sierra de la Venta, 18 km W of Guadalajara, Villarreal de Puga 531 (eNcB). Belize: Augustine, 16°34’N, 88°54'W, Hunt 438 (BM, Us); El Cayo dist.. m S of Georgeville, Liesner & Dwyer 1612 (mo, Ny); El Cayo dist., San Agustin. Lundell 6792 (F, NY, US). Honduras: Las Jagnas, Minas de Oro, Comayagua dist., Edwards P-180 (a), El Paraiso, rd. from Zamoras to Giiinope, Webster et al. 11988 (F, GH, MO. us); dept. Morazan, Rd. Valle de Angeles near San Antonio, Molina 389 (F, GH, MEXU. MO, UC, US); dept. El Paraiso, between Galeras and Las Casitas, Molina 10765 (F, us): dept. Comayagua, La Piramide, Molina 14393 (Ff); dept. Morazan, San Antonio de Oriente, Rodriguez 3133 (F, GH, NY); dept. El Paraiso, NW of Giiinope, Standley et al. 2047 (F, NY, US); dept. Morazan, vic. of Agua Amarilla, Standley 12297 (Fr); dept. E] Paraiso, above Galeras, Williams & Molina 10246 (F, GH); dept. Comayagua, 5 km N of Siguatepeque, Williams & Molina 18128 (F, Gu, US). Agarista mexicana is closely allied to A. sleumeri and A. populifolia; see species are characterized by ovate leaves with acuminate apices and ro c to cuneate bases, and by short inflorescences of white flowers. ee mex- icana can be differentiated from both taxa by its pubescent ovary, from 4. populifolia by its corky, prominently furrowed bark, and from A. s/eumeri by its pubescent calyx and pedicels and its smaller corollas and leaves. Agarista mexicana is a widely distributed taxon (see Map 3) that is extremely variable in its unicellular indumentum. Populations of this species are here divided into two weakly differentiated geographic varieties on the basis of the density of unicellular hairs on the abaxial leaf surface. However, some popu- lations are variable in leaf pubescence, and intermediate specimens are known, mostly in the state of México. Representative intermediate material includes: Mexico. CHiApas: near Fenia, Purpus 294 (us); E of Monserrate, Purpus 10156 (A, F). Purpus 10518 (a, NY). México: dist. Temascaltepec, Ypericones, Hinton 3899 (GH, NY. us); 5 km SW of Nanchititla, Tejupilco, Medrano et al. 5030 (mMexu);, Puerto del Em- bocadera, 7 km W of Luvianos, municipio de Tejupilco, Rzedowski 22122 (ENCB, MEXU), La Cienaga, 4 km S of Sultepec, Rzedowski 30392 (ENCB). Agarista mexicana var. mexicana has been illustrated by Standley and Wil- liams (1952, fig. 30). 7. Agarista sleumeri Judd, sp. nov. FiGure 3, a. Frutex vel arbor ad 6(—20) m alta. Ramuli hornotini glabri cum medulla septata. Folia ovata, 4—-11.5(—13) cm longa, 1.6—4.5(—5.3) cm lata, coriacea, ad apicem acuminata, ad basin cuneata vel rotundata; margo + planiuscula, in- tegra; pagina abaxialis glabra; petioli 6.5—13 mm longi. Inflorescentiae axillares, racemosae, ad 0.5—2.5(—4) cm longae; axis glaber vel leviter pubescens. Pedicelli glabri, 3.5-8.5 mm longi. Flores 5-merus. Calyx lobis |.2—2.6(-3) mm longis, 0.7-2 mm latis, abaxialiter glabris. Corolla cylindrica, 8.5—13 mm longa, 3-- 7.5 mm lata, alba, abaxialiter glabra. Filamenta 5—7.5 mm longa; antherae | .3-- 1.7 mm longae. Ovarium glabrum. Capsula subglobosa vel brevissime ovoidea, 3-4.5 mm longa, 5-8 mm lata, placentis subapicalibus. Semina 1.5-2.4 mm longa. 294 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Ficure 3. a, Agarista sleumeri (Koch & Garcia 76103). b, A. oleifolia var. oleifolia (Glaziou 15171). c, A. coriifolia var. bradei (Irwin et al. 28486): note round-based | pees with conspicuous glandular dots extending along midvein on abaxial surface. d, 4. co- rilfolia var. coriifolia (Duarte 2044): note cordate-based leaves with glandular fe less well developed. Scale = 1984] JUDD, AGARISTA 295 Shrub or tree to 6(—20) m tall, with corky, longitudinally furrowed bark. Twigs glabrous, with clearly chambered pith. Buds to ca. 1.2 mm long. Leaves alternate (to rarely subopposite at few nodes); petiole 6.5—13 mm long; blade revolute in bud, ovate, 4-11.5(-13) by 1.6—4.5(-5.3) cm, flat, coriaceous, the apex acuminate, the base cuneate to rounded, the margin entire (slightly un- dulate), plane to very slightly revolute near base, the adaxial surface glabrous to very slightly pubescent near extreme proximal portion of midvein, the abax- ial surface glabrous, with few to several inconspicuous glandular dots along midvein. Inflorescences (fasciclelike) axillary racemes to 0.5—2.5(—4) cm long, axis essentially glabrous to sparsely pubescent. Pedicels 3.5—8.5 mm long, gla- brous; bracteoles 2, alternate, nearly basal to within lower 4 of pedicel, tri- angular to narrowly so, to ca. 0.8 mm long; bracts to 1.3 mm long. Calyx lobes triangular with acuminate apices, 1.2-2.6(-3) by 0.7-2 mm, abaxial surface glabrous; corolla cylindrical, 8.5-13 by 3-7.5 mm, white, abaxially glabrous; filaments 5—7.5 mm long, anthers 1.3-1.7 mm long; ovary glabrous. Capsules subglobose to short-ovoid, 3—4.5 by 5-8 mm, placentae subapical; seeds |.5- 2.4 mm long. Type. Mexico, Hidalgo, municipio de Tenango de Doria, 30 km NE of Tulan- cingo, rd. between Metepec and Tenango de Doria, 19 km N of Metepec, 22 May 1976, S. D. Koch & J. Garcia P. 76103 (holotype, ENCB!; isotypes, MEXU!, MO!). DISTRIBUTION AND ECOLOGY. Mexico, limited to small area of Hidalgo, Vera- cruz, and Puebla in Sierra Madre Oriental (MAP 3). Pinus and/or Quercus forests, Pinus and Liguidambar forest, mixed forest of Pinus, Quercus, Alnus, or Cupressus, and Lyonia and Cupressus forest; 1220-2250 m alt. Flowering April and May (June). REPRESENTATIVE SPECIMENS. Mexico. HipALco: 2 km SW of Tutotepec, Gimante 608 (ENCB); municipio de Tenango de Doria, El Estribo, Gimante 950 (ENCB, MEXU, Wis), NE of Honey, Hernandez 767 (MEXU); municipio de Tenango de Doria, 25 km NE of Tu- ango, Sharp 45353 (mexu); | km from Tlaxpanaloya, Naupan, Vela 455 (ENcB). VER- ACRUz: La Capilla, 3 km from Huayacocotla, Calzada & Horritz 2628 (F, MEXU, NY); SW entrance to Huayacocotla on rd. from Palo Bendito, Diggs & Nee 2957 (F, FLAs); El Paraje, Huayacocotla, Hernandez & Cedillo 817 (F, GH, MEXU, MO); Municipio Huaya- cocotla, Hernandez 1150 (mexu); between La Cruz del Ataque and Zacualpan, Vela 618 (ENCB). Agarista sleumeri is most closely related to 4. populifolia and A. mexicana, differing from both in its abaxially glabrous calyx and glabrous pedicel (vs. very sparsely to densely pubescent calyx and pedicel) and its larger corolla. It can also be distinguished from the latter species by its glabrous (vs. very sparsely to densely pubescent ovary, its clearly septate pith, and its usually larger leaves. These three species foie a well-marked subgroup of the genus and can be separated from the South American species by their ovate leaves with cuneate 296 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 a ne ae oo eee 3 FiGure 4. Agarista populifolia (Judd oo a, segment of twig, 5; b, sees section of twig, showing chambered pith, , cross section of olde er stem, showing hollow central portion, x 1; d, margin of leaf. x 10; e, multiseriate-stalked, gland- ee hair of Boe x 130; f, unicellular hairs of pedicel, x 130; g, flower, x 5; h, x 1031 > 1, anther, x 25: j, capsule and portion pedicel, x 5; k, columella with ee x 6.5: ei seed with elongated testa cells, x 10. 1984] JUDD, AGARISTA 207 to rounded bases and acuminate apices, their white flowers, and their short racemes. The three species are geographically isolated (Maps 3, 4). It is a pleasure to name this distinctive species for Dr. H. Sleumer, who has done so much to increase our taxonomic understanding of Agarista and various other ericad groups. 8. Agarista populifolia (Lam.) Judd, Jour. Arnold Arb. 60: 195. 1979. FIGURE 4. gees populifolia eee Encycl. Méth. Bot. 1: 159. 1783. Andromeda laurina or.-Am. 1: 253. 1803, nomen superfl. Lyonia populifolia (Lam.) K. Sean Deadral 2: 123. ie Leucothoé populifolia (Lam.) Dippel, Handb. Laubh. 1: 356. 1889. Tyre: United States, South Carolina, Fraser s.n. (holotype, P-LA; photo of holotype, GH!). pret lucida Jacq. Collect. 1: 95. es non Andromeda lucida Lam. Encycl. 1: 157. 3 (= Lyonia lucida (Lam.) K. Koch). Pree reticulata Walter, FI. Carolin. ee 1788. Type: not seen Andromeda acuminata Aiton, Hortus Kew. 2: 70. 1789. Andromeda acuminata Lodd. ex C. F. Ludwig, Neue Wilde Baumz. 4. 1783, nomen nudum. Leucothoé acuminata (Aiton) G. pes Gen. Syst. 3: 832. 1834. Tyre: cultivated, “introduced by Mr. John Cree”’ (not ). Andromeda formossina Bartram, Travels N. S. Carol. 5: 24, 172, 303. 1791, nomen nudum Shrub or small tree to 7 m tall, with shallowly furrowed bark. Twigs with or without scattered gland-headed hairs, otherwise glabrous to sparsely pu- bescent, with clearly chambered pith. Buds to ca. | mm long. Leaves alternate; petiole 3-12 mm long; blade revolute in bud, ovate, 2.6-9(-11.2) by 0.9-4(-5) cm, flat, coriaceous, the apex acuminate (to eae pane the base harrow cuneate to rounded, the margin entire to obscurely o when present, each associated with gland-headed hair) Goes blac to a slightly revolute near base, the adaxial surface with or without scattered gland- headed hairs, otherwise sparsely pubescent on midvein, especially proximal portion, the abaxial surface with or without gland-headed hairs along midvein (and proximal portion of lamina), otherwise glabrous or with very few uni- cellular hairs at base of midvein, with few to several inconspicuous glandular dots along midvein. Inflorescences (fasciclelike) axillary racemes to 1—2.5(-3.5) cm long, the axis with or without scattered gland-headed hairs, otherwise sparsely to densely pubescent. Pedicels 5.5-11.5 mm long, with or without gland-headed hairs, otherwise sparsely to moderately pubescent; bracteoles 2 (rarely 3), alternate, nearly basal to within lower 3 (rarely to midpoint) of pedicel, triangular or narrowly so, to ca. 1.2 mm long, very rarely subtending an axillary flower; bracts to 1.2 mm long. Calyx lobes triangular with acuminate apices, 0.9-2 by 0.6-1.5 mm, the abaxial surface with or without gland-headed hairs, otherwise sparsely to moderately pubescent; corolla cylindrical, 6-9.5 by 3-5 mm, white, abaxial surface glabrous; filaments 4-5.5 mm long, anthers 1.1-1.4 mm long; ovary glabrous. Capsules subglobose to short-ovoid, 3-4 by 4.5-6.5 mm, placentae subapical; seeds 1.4-2 mm long. 27 = 24 (Wood, 1961). DISTRIBUTION AND ECOLOGY. Southeastern U. S. Atlantic Coastal Plain from southern South Carolina to eastern Florida (Map 4). Acidic swamps in pine 298 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 84 83 Map 4. cality). Distribution of Agarista populifolia (open circle = unspecified Georgia lo- 1984] JUDD, AGARISTA 299 flatwoods, mixed hydric hammocks (hardwood swamps) of Fraxinus, Quercus, Nyssa, Sabal, Acer, Liquidambar, Persea, Gordonia, Carpinus, and Pinus, riv- erine swamps, swamp forests of Taxodium or Chamaecyparis (along with various hardwoods), bayheads and lake margins, frequently along streams or in ravines; near sea level to 55 m alt. Flowering chiefly late March through early COMMON NAMES. Pipe-plant, pipe-stem wood. REPRESENTATIVE SPECIMENS. United States. FLoripa. Clay Co.: ravine at Goldhead Branch St. Park, NE of Keystone Heights, Arnold s.n., 5 May 1940 (FLas); Rte. 21, 4.8 mi N of jet. with Rte. 16, along Mill Creek, $22, TSS, R24E, Judd 2541 (FLAs); 5 mi W of Penny Farms, $15, T6S, R24E, Ward 5521] (FLAs). Duval Co.: near Jacksonville, Curtiss 1692 (A, G, GH, K, M, MO, NY, UC, us). Lake Co.: Alexander Springs, Ford & West s.n., 11 July 1958 (FLAS, UNC). Marion Co.: just N of Orange Springs, near jct. of Orange Creek and Rte. 21, $25, T11S, "ROSE, Judd 2609 (FLAS, FSU, GH, MO, S, UNC); Juniper Springs, Murrill s. n., 5 May 1940 (FLas, Mo). Nassau Co.: near Callahan, £. J. Palmer 38293 (a, Mo, s). Orange Co.: Rock Springs, $15, T20S, R28E, Moore s.n., 1 April 1961 a May-June 1875 (mo, ny). GeorGiA: without definite locality, Le Conte s.n. (Mo); Feay s.n. (NY). SOUTH CAROLINA. Beaufort Co.: Bluffton, Mellichamp s.n., 1876 (GH, NY). Hampton Co.: Garnett, Lawton s.n., 28 Sept. 1931 (Ncu). Jasper Co.: Elliott (1817) reported it from near the “Black Swamp, on the road from Coosawhatchie to the Sisters ry.” Agarista populifolia, the northernmost-ranging species of the genus, is very closely related to A. sleumeri. The two species are distinguished by calyx and pedicel indumentum, corolla size, and bark characters, and they are geograph- ically separated, with A. populifolia occurring on the Atlantic Coastal Plain from South Carolina to Florida and A. slewmeri in the Sierra Madre Oriental of Hidalgo, Veracruz, and Puebla. Agarista populifolia is also allied with A. mexicana, from which it is easily distinguished by its glabrous (vs. pubescent) ovary, its shallowly (vs. corky and prominently) furrowed bark, and its more consistently septate pith. It is puzzling that 4. populifolia has consistently been misclassified despite the many characters linking it to its South and Central American relatives (see TABLE 1). Agarista populifolia is rather variable in glandular indumentum and in the size and margin of the leaves. Leaves of some plants are sharply (to obscurely) serrate, while those of others are entire; the two forms are commonly found intermixed within the populations. In addition, some plants have gland-headed hairs on the inflorescences and frequently also on the stems and/or leaves, while others completely lack them. Most populations contain plants with such hairs and plants without them. This species is locally common in eastern Florida (Map 4) from Osceola County to Nassau County but is apparently extremely rare in Georgia and South Carolina. In fact, the presence of the taxon in Georgia is somewhat questionable since no exact localities are known, although Bartram (1791) 300 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 65 reported it growing along the St. Marys River. The species is occasionally cultivated (Ingram, 1961, fig. 26; Melvin, 1981). Andromeda serratifolia (see DC. Prodr. 7: 602, 609. 1839) and 4. serrata (see ibid. 602) are horticultural names (of no nomenclatural standing) that were occasionally used for Agarista populifolia in the late eighteenth or early nine- teenth century. 9. Agarista coriifolia (Thunb.) J. D. Hooker ex Niedenzu, Bot. Jahrb. 11: 236. Shrub (to tree) to 2(-5) m tall. Twigs glabrous to moderately pubescent (glaucous), with nonchambered to clearly chambered pith. Buds to ca. 1.8 mm long. Leaves alternate to subopposite or nearly 3-whorled at some nodes; petiole 2-18 mm long; blade revolute in bud, ovate to elliptic or oblong, 1.8-10.5 by 0.6-3.5(—5) cm, + flat, very coriaceous, the apex acute- to rounded-mucronate to acuminate, the base cordate to cuneate, the margin entire, plane (slightly revolute near base, rarely slightly undulate), the adaxial surface glabrous to sparsely pubescent on midvein, especially proximal portion, the abaxial surface glabrous to very sparsely pubescent along proximal portion of midvein, with glandular dots absent, or few to many and inconspicuous to conspicuous along midvein. Inflorescences axillary or terminal racemes or panicles, to (2.5—)4.5- 28 cm long, axis glabrous to densely pubescent. Pedicels 2-14 mm long, gla- brous to densely pubescent; bracteoles 2 (rarely to 4), alternate to opposite, from nearly basal to near middle of pedicel, triangular to narrowly so, to ca. 1.6 mm long; bracts to 4 mm long. Calyx lobes tnangular with acuminate apices, 1—2.7 by 0.6-2.2 mm, abaxial surface glabrous to sparsely (moderately) pubescent; corolla cylindrical to urceolate-cylindrical, 6.5-11 by 5 white or greenish white to red, abaxially glabrous (rarely with very few uni- cellular hairs near apex); filaments 4-6 mm long, anthers |.3-—2.3 mm long; ovary glabrous to moderately pubescent. Capsules short-ovoid to subglobose, 3-6 by 4.5-8 mm, placentae subapical (rarely + central); seeds 1-2.6 mm long. DistRiBUTION. Eastern Brazil, Bahia, Minas Gerais, and Estado do Rio. KEY TO THE VARIETIES OF AGARISTA CORIIFOLIA — Leaves usually ovate (to + elliptic), base usually cordate to rounded, abaxial surface with glandular dots lacking to limited to aie half of blade; calyx lobes abaxially glabrous to sparsely (moderately) pubescent. .................. 9a. var. cortifolia. Leaves + oblong to slightly elliptic ay ovate), base usually cuneate to rounded, abaxial surface with glandular dots usually extending along midvein to near apex of blade; calyx lobes abaxially glabrous. ................0..00.0005. b. var. bradei. — 9a. Agarista coriifolia (Thunb.) J. D. Hooker ex Niedenzu var. coriifolia FiGure 3, d. Andromeda coriifolia Thunb. Pl. Brasil. Decas 1, p. 9. 1817. Leucothoé coriifolia (Thunb.) DC. Prodr. 7: 605. 1839. Type: Brazil, aie Gerais, Villa Rica, Frevreiss s.n. (holotype, UPS). Leucothoé neriifolia Cham. & Schldl. Linnaea 1: 522. 1826. Agarista neriifolia (Cham. & Schldl.) G. Don, Gen. Syst. 3: 838. 1834. Leucothoé laxiflora Meissner var. sellowii 1984] JUDD, AGARISTA 301 Meissner in Martius, Fl. Brasil. 7: 157. 1863 (= var. /axiflora), nomen superfl. Tyre: Brazil, Minas Gerais, without definite locality, Se/low s.n. psu B (destroyed); fragment of holotype, Ny; photos of holotype, F!, G!, GH!; isotypes, E!, K!). Andromeda crassifolia Pohl, Pl. Brasil. 2: 34. 1828/29, non Nees, saa 297. 1821. Agarista pohlii G. Don, Gen. Syst. 3: 837. 1834, nomen novum. Leucothoé crassifolia (Pohl) DC. Prodr. 7: 605. 1839. Leucothoé crassifolia (Pohl) DC. var. subevenia Meissner in Martius, Fl. Brasil. 7: 158. ¢. 59, fig. J. 1863 (= var. crassifolia), nomen superfl. Leucothoé pohlii (G. Don) Sleumer, Bot. Jahrb. 78: 463. 1959. Type: Brazil, Minas Gerais, Dumienine, Pico Itambé, Nov. 1821, Poh! 3506 (holotype, w; frag- ment of holotype, Ny Leucothoé laxiflora oe var. martii Meissner in Martius, Fl. Brasil. 7: 157. 1863. Type: Brazil, Estado do Rio, near Paraiba, Martius 1118 (holotype, m!; fragment of holotype, ny!; photos of holotype, F!, GH!; isotypes, M!). Leucothoé laxiflora Meissner var. subcordata Meissner in Martius, ibid. Tyre: not Teicainoe laxiflora Meissner var. hookeriana Meissner in Martius, ibid. 158. Type: cultivated in ea in garden of Mr. Cunningham (see Hooker & Smith, 1851, t. 4593) (holotype, k!). Leucothoé ears (Pohl) DC. var. subreticulata Meissner in Martius, ibid. Type: Brazil, ee Gerais, Itambé, Aug. 1840, Martius 836 (holotype, m!; fragment of holotype, Ny!; isotypes, BR, G!, K!, am ; photos of isotype, F!, G Pa ees se (Pohl) DC. var. reticu ilat a Meissner in Martius ibid. Type: Brazil, Minas wie rais, without definite locality, Claussen 535 (holotype, BR; fragment of holotype, NyY!). Leucothoé en (Pohl) DC. var. macrophylla Meissner in Martius, ibid. Type: Brazil, Serra da Paraca, fl. Paraiba, Martius s.n. (holotype, m!; photos of holotype, F!, GH!). Shrub (or tree) to 2(—5) m tall. Twigs glabrous to moderately pubescent. Leaves with petiole 2-18 mm long; blade ovate (to + elliptic), 1.8-10.5 by (0.6—-)1-3.5(—5) cm, base usually cordate to rounded, abaxial surface with glan- dular dots absent to limited to proximal half. Inflorescence axis to 6-28 cm long, glabrous to densely pubescent. Pedicels glabrous to densely pubescent. Calyx lobes abaxially glabrous to sparsely (or moderately) pubescent; corolla 6.5-10.5 mm long, red to white; ovary glabrous to sparsely pubescent. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, from Bahia to southern Mi- nas Gerais and adjacent Estado do Rio (Map 5). Open rocky and/or grassy campo, open scrub, thickets and forests, frequently along streams; usually in sandy soil, with sandstone or manganese outcrops; 500-1950 m alt. Flowering chiefly June through September (to January or February). REPRESENTATIVE SPECIMENS. Brazil. BAntA: Lengéis, Duarte 9337 (L); Serra do Sincora, by Rio Cumbuca, ca. 3 km S of Mucugé, Harley et al. 15911 (£, L, MO, Ny, us); 10 km S of Andarai on rd. to Mucugé, Harley 18758 (Ny); 3 km S of Mucugé, Mori et al. 12578 (Ny); between Len¢dis and Itaberaba, Pereira 2040 (1); Serra do Sincora, Ule 7336 (HBG, L). Minas Gerais: E slopes of Pico do Itambé, Anderson et al. 35887 (c, F, L, NY, US); Serra do Taquaril, Belo Horizonte, Barreto 528 (F); Serra do Cipé, km 131, Palacio, Santa Luzia, Barreto 9253 (Fr); Cachoeira do Campo, Claussen 54 (a); Palmeira, Rio Carandai, Duarte 4314 (L); Caraca, Emygdio et al. 3525 (Ny); Furnas, Alvinépolis, Emyedio et al. 3588 (Ny); Ribeirao oe Capidalas, Emygdio 3623 (Ny); Diamantina, Gardner eae (E, G, GH, K, NY, P, US); Serra do Palmital, near Sdo Bartholomeu, Glaziou 15172 (c, p); Serra do ee Gomes 2846 (F); Pico Itambé, Sao Antonio do Itambé, Hisicek 30123 (c, L, MO, UC); Serra do Caraga, Irwin et al. 29067 (L, Mo, 302 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Map 5. ey of se ee coriifolia var. cortifolia (circles), A. coriifolia - bradei (solid squares), A. a (open squares), A. subrotunda (open triangles), chapadensis (solid pices) a eee var. revoluta (dots), and A. revoluta var. ne (star). NY), St.-Hilaire s.n., 1816-1821 (p); Serra do Cip6, Mato Dentro, Macedo 3757 (Ny, s, us); Serro Frio, Martius s.n. (mM); Serra do Curral d’El Rey, Reinhardt s.n., Sept. 1855 (c); Serra d’Uro Branco, St.-Hilaire 234 (p); Itambé, St.-Hilaire 395 (p), Serra do Ouro Préto, Ule s.n., Feb. 1892 (HBG); Casa Branca, Ouro Préto, Williams 8130 (GH). 9b. iat coriifolia (Thunb.) J. D. Hooker ex Niedenzu var. bradei (Sleu- er) Judd, comb. et stat. nov IGURE 3, ¢c. Leucothoé bradei Sleumer, Notizbl. Bot. Gart. Berlin 13: 213. 1936. Type: Brazil, Minas Gerais, Diamantina, June 1934, Brade 13614 (holotype, RB; fragment of Rei LIL; photos of isotype, B (destroyed), s!). Shrub to 2 m tall. Twigs glabrous to very sparsely pubescent. Leaves with petiole 4-11 mm long; blade + oblong to slightly elliptic (slightly ovate), (2.4-)3-7.5 by 0.6—2.5(—3) cm, base usually cuneate to rounded, abaxial surface with glandular dots usually extending along midvein from base to near apex. Inflorescence axis to (2.5—)4.5—15(-20) cm long, glabrous to sparsely pubescent. 1984] JUDD, AGARISTA 303 Pedicels glabrous to sparsely pubescent. Calyx lobes abaxially glabrous, corolla 8-11 mm long, red; ovary glabrous to moderately pubescent. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, Minas Gerais (Map 5). Rocky or grassy campo, shrubby cerrado, gallery forest, sometimes on termite mounds; ca. 1200-1410(?) m alt. Flowering September and October. REPRESENTATIVE SPECIMENS. Brazil. Minas Gerais: Serra do Espinhaco, 13 km SW of Rio Jequiti and Mendanha, on rd. to Diamantina, Anderson 8992 (Cc, F, L, MO, NY); between Diamantina and Guinda, Archer 4103 (A, Ny); Diamantina, Duarte 8927 (L); Guinda, Diamantina, Hatschbach 27371 (c, L, NY, 8, UC, US); 27 km SW of Diamantina on rd. to Gouveia, Irwin et al. 22043 (F, L, Ny); 3 km N of Sao Joao da Chapada, rd. to Campo do Sampaio, Jrwin 28486 (F, L, Ny); Chapada, St.-Hilaire, Cat. B’, n. 2048 (P). Agarista coriifolia is probably closely related to A. pulchraand A. subrotunda. It can be separated from the former by its larger, narrower leaves with a length/ width quotient of (1.5-)1.8-6 (vs. 0.9-1.7) and its glabrous to densely pubescent inflorescence axes, and from the latter by its abaxially glabrous leaves. This species may also be confused with A. pulchella var. pulchella or A. oleifolia (both varieties), but these taxa have more or less central placentae and usually less coriaceous leaves that always lack conspicuous abaxial glands (see key for other distinguishing features). Agarista coriifolia is extremely variable in unicellular indumentum, shape and size of the leaves, development of abaxial laminar glands, and color of the corolla. This variation has led to the description of numerous species and varieties (e.g., A. neriifolia, A. pohlii, A. bradei, or the various varieties of A. laxiflora) that cannot be maintained when the total pattern of variation is analyzed. Plants with glabrous inflorescence axes and often large leaves lacking abaxial glandular dots along the midvein have often been considered as A. neriifolia or A. coriifolia sensu stricto, while individuals with pubescent inflo- rescence axes, and small to large leaves with few to many, conspicuous to inconspicuous abaxial glands along the midvein have been treated as 4. pohlii. However, these characters are not consistently correlated, and intermediate specimens—e.g., Harley 15911 (E, L, MO, Ny, US), Harley 18758 (Ny), St.- Hilaire s.n., 1816-1821 (p), St.-Hilaire 234 (pr), Claussen 5 (Bm), Claussen 214 (c), Barreto 528 (F), Williams 8130 (Gu), and Pereira 2040 (L)—are common. In addition, both morphological extremes, along with intermediates, grow in the same geographic regions. Thus, these plants are here considered as repre- sentatives of a single variable species. Yet specimens previously referred to A. bradei (occurring in Minas Gerais, from Diamantina to Chapada in the Serra do Espinhaco) are distinctive due to their leaves that are more or less oblong to slightly elliptic, with cuneate to rounded bases and numerous very conspic- uous glandular dots along the midvein abaxially. These populations are here maintained as a weakly delimited variety of A. coriifolia. The remaining more variable populations are all treated under var. coriifolia due to the lack of any internal discontinuities in the pattern of variation. The two varieties are, at least in part, geographically separate (Map 5), but both may occur together in the Diamantina region, and the factors isolating them are in need of field investigation. 304 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 10. Agarista pulchra (Cham. & Schldl.) G. Don, Gen. Syst. 3: 837. 1834. Andromeda pulchra Cham. & Schldl. Linnaea 1: 521. 1826. Leucothoé pulchra (Cham. & Schidl.) DC. Prodr. 7: 604. 1839. Andromeda sellowii Steudel, Nomencl. Bot. 2: 89. 1841, nomen superfl. Type: Brazil, without caer Sellow s.n. (holotype, B (destroyed); fragment of holotype, Ny; isotypes, E Gaylussacia leptobotrys DC. Prodr. 7: 560. 1839. Ty PE: el Minas Gerais, Serra ae re Nov. 1834, Lund s.n. (holotype, G-pc; fragment of holotype, Ny; iso- Gl), ie pulchra (Cham. & Schldl.) DC. var. parvifolia Meissner in Martius, Fl. rasil. 7: 161. 1863. Type: Brazil, Minas Gerais, without definite locality, 1843, Claussen 159 (lectotype (selected by Sleumer, 1959), Br; isolectotypes, G!, Ny!). Shrub to ca. | m tall. Twigs glabrous to very sparsely pubescent, with non- septate pith. Buds to ca. 1.3 mm long. Leaves alternate; petiole 1-5 mm long: blade revolute in bud, ovate to elliptic, 1.3-3.5(-4.3) by 0.8-—2(-2.5) cm, + flat, usually very coriaceous, the apex acute- to rounded- or retuse-mucronate, the base cordate to rounded, the margin entire, plane, the adaxial surface glabrous to sparsely pubescent on proximal portion of midvein, the abaxial surface glabrous, usually with few to several inconspicuous to conspicuous glandular dots along midvein. Inflorescences axillary racemes to 4-12 cm long, axis glabrous to very sparsely (moderately) pubescent. Pedicels 3.5—7(-8) mm long, glabrous to sparsely (rarely moderately) pubescent; bracteoles 2, alternate to subopposite, from nearly basal to within lower '4 (to near middle) of pedicel, triangular to narrowly so, to ca. 1.2 mm long; bracts to 1.7 mm long. Calyx lobes triangular with acuminate apices, 1.2-2 by 0.6-1.3 mm, abaxial surface glabrous (to rarely sparsely to moderately pubescent); corolla cylindrical, 6- 8.5(-9) by 2.5-4 mm, white (to reddish), abaxially glabrous; filaments 3.5—5 mm long, anthers 1.1-1.3 mm long; ovary glabrous (rarely sparsely to mod- erately pubescent). Capsules subglobose to short-ovoid, 3.5—-4.5 by 5-7 mm, placentae subapical (to occasionally + central); seeds 1-1.5 mm long. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, small region of south-central Minas Gerais (MAP 5). Open campo and slopes, low, dense woodland and thickets; iron-rich soil. Flowering September through January. REPRESENTATIVE SPECIMENS. Brazil. MINAS GERAIS: Camarinhas, Serra do Ouro Préto, Barreto 9065 (F); Morro de Santa Anna, Ouro Préto, Barreto 9070 (F); Serra da Piedade, Damazio 1240 (G), Glaziou 20390 (c, G, Kk), Riedel 2666 (G, L, NY); Serra de José d’E] Rey and Ouro Préto, Campo de Sao Sebastiaéo, Glaziou 15173 (c), Caeté, Serra da Piedade, Janna & Strang 1975/1486 (1); Serra da Piedade, ca. 40 km E of Belo Horizonte, near BR-31, /rwin et al. 30468 (Ny, us); Serra de Ouro Branco, St.-Hilaire s.n., 1816- 1821 (ny, P) Agarista pulchra is apparently closely related to A. coriifolia and A. subro- tunda. It can be distinguished from the former by its usually glabrous (vs. glabrous to densely pubescent) inflorescence axis and its usually smaller, broad- er leaves with a length/width quotient of 0.9-1.7 (vs. ca. 1.8-6), and from the latter by its glabrous abaxial leaf surface and corolla. The factors isolating these taxa are in need of field study. See illustration in Hooker (1847, t. 4313 Several specimens—for example, St.-Hilaire s.n., 1816-1821 —are unusual 1984) JUDD, AGARISTA 305 because of their rather large leaves and their capsules with central placentae. These plants are tentatively placed here, although they also show similarities to Agarista coriifolia. 11. Agarista subrotunda (Pohl) G. Don, Gen. Syst. 3: 837. 1834. Andromeda subrotunda Pohl, Pl. Brasil. 2: 32. t. 127. 1828/29. Leucothoé subrotunda (Pohl) DC. Prodr. 7: 603. 1839. Type: Brazil, Minas Gerais, Sdo Joao d’El Rey, Oct 1819, Pohl s.n. (not seen). Andromeda pistrix Cham. Linnaea 8: 508. 1833. Agarista pistrix (Cham.) G. Don, Gen. Syst. 3: 838. 1834. Leucothoé pistrix (Cham.) DC. Prodr. 7: 604. 1839. Leu- cothoé subrotunda (Pohl) DC. var. pistrix (Cham.) Meissner in Martius, Fl. Brasil. 7: 161. 1863. Type: Brazil, Minas Gerais, Serra do Lenheiro, Sellow s.n. (holotype, B (destroyed); fragment of holotype, Ny). Shrub to ca. 2.5 m tall. Twigs with or without scattered gland-headed hairs, otherwise densely pubescent, with nonseptate pith. Buds to ca. 1.2 mm long. Leaves alternate; petiole 2-4.5 mm long; blade revolute in bud, ovate to elliptic or nearly orbicular, 1-3.5(-4) by 0.7-2.5(-3.7) cm, flat to slightly abaxially curved, very coriaceous, the apex acute- to rounded- or retuse-mucronate, the base cordate, the margin entire (ciliate due to gland-headed hairs), + plane, the adaxial surface sparsely pubescent on lamina and midvein, often glabres- cent, the abaxial surface with or without gland-headed hairs on midvein, oth- erwise moderately to densely pubescent on lamina and sparsely to densely pubescent on midvein, with few to several inconspicuous to conspicuous glan- dular dots along midvein (although these often obscured by dense pubescence). Inflorescences axillary racemes to 4-7 cm long, the axis with or without scat- tered gland-headed hairs, otherwise densely pubescent. Pedicels 5—12.5 mm long, with or without gland-headed hairs, otherwise densely pubescent, brac- teoles 2, alternate to subopposite, from nearly basal to near middle of pedicel, narrowly triangular, to ca. 1.3 mm long; bracts to 2 mm long. Calyx lobes triangular to ovate, with acuminate apices, 1.5-2.5 by 1-2 mm, the abaxial surface sometimes with few gland-headed hairs, otherwise densely pubescent; corolla cylindrical, 6.5-9 by 3—4.5 mm, red, abaxially sparsely to densely pu- bescent; filaments 3.5-4.5 mm long, anthers 1.5—2 mm long; ovary densely pubescent. Capsules short-ovoid, 4-5 by 6-7 mm, placentae subapical; seeds 1.3-2 mm long (probably also larger). DISTRIBUTION. Brazil, southern Minas Gerais, region surrounding Sao Joao d’El Rey (Map 5). Circa 1065-1675 m alt. Flowering July and August. REPRESENTATIVE SPECIMENS. Brazil. MINAS GERAIS: Serra de Tiradentes, Barreto 4787 (L); Serra do Lenheiro, Glaziou 17110 (c, P); SAo Jodo dEl Rey, Sellow s.n. (E, G, K) Stephan s.n., Aug. 1876 (k). Agarista subrotunda is likely closely allied with A. pulchra and A. coriifolia. All of these species have thickly coriaceous, at least occasionally conspicuously gland-dotted leaves, and usually subapical placentae. This species is easily separated from both A. coriifolia and A. pulchra by its conspicuous abaxially pubescent leaves and corollas. Agarista chapadensis has similar abaxially pu- 306 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 bescent leaves and flowers and may also be closely related; it can be distin- guished from A. subrotunda by leaf-base shape (cuneate to rounded vs. cordate) and pedicel length. Agarista paraguayensis, A. mexicana var. pinetorum, and A. revoluta var. velutina also have abaxially pubescent leaves, but these taxa differ in many other characters and are not closely related to A. subrotunda. Agarista subrotunda appears to be geographically separated from all of the above-mentioned species except 4. pulchra (Maps 3, 5, and 6). The taxon has been poorly collected and is in need of field inyestioation. 12. Agarista chapadensis (Kinoshita-Gouvéa) Judd, comb. nov. Leucothoé chapadensis Kinoshita-Gouvéa, Rev. Brasil. Bot. 4: 127. fig. 2. 1981. Type: Goids, Chapada dos Veadeiros, 20 km N of Alto Paraiso, 1600 m alt., 6 March 1983, Anderson et al. 6502 (holotype, UNB; isotype, NY!). Shrub or small tree to ca. 3 m tall. Twigs moderately to densely pubescent, with nonseptate pith. Buds to ca. | mm long. Leaves alternate; petiole 2-6 mm long; blade revolute in bud, oblong or elliptic to slightly ovate, 2.4-6 by 0.7- 2.2. cm, flat, coriaceous, the apex acute- to rounded-mucronate, the base cuneate to rounded, the margin entire, plane, the adaxial surface sparsely pubescent on midvein, the abaxial surface densely pubescent on lamina and midvein (hairs frequently ferrugineous, at least when young) (with few + inconspicuous glan- dular dots along midvein, these obscured by dense pubescence). Inflorescences axillary racemes to 4-6 cm long, axis densely pubescent. Pedicels 3-7 mm long, sparsely to densely pubescent; bracteoles 2, alternate to subopposite, from nearly basal to near middle of pedicel, narrowly triangular, to ca. 1 mm long; bracts to ca. 1.3 mm long. Calyx lobes triangular with acuminate apices, 1-2 by 0.7-1.5 mm, abaxial surface moderately to densely pubescent; corolla cy- lindrical, 6-8 by 3-4.5 mm, white, abaxially moderately to densely pubescent; filaments 4-6 mm long, anthers 1|.1-1.4 mm long; ovary densely pubescent. Capsules subglobose to short-ovoid, 4.5-6 by 6-7 mm, placentae subapical: seeds |.5-3 mm long. DISTRIBUTION AND ECOLOGY. Central Brazil, Goias and Bahia (Map 5). Sandy or rocky campo, rocky (sandstone) open cerrado, moist woods along streams; ca. 1000-1600 m alt. Flowering February and March. RESEN E SPECIMENS. Brazil. BAHIA: 4 km N of Rio de Contas, Mori et al. 12412 (is). Goras: (Chapada dos Veadeiros, ca. 10 km W of Veadeiros, Irwin et al. 12862 (FLAS, L, NY). Agarista chapadensis is probably allied with A. subrotunda, however, both taxa have been poorly collected and their relationship will remain somewhat obscure until additional material has been collected. The two taxa are distinc- tive due to their abaxially conspicuously pubescent leaves. Among the South American species of Agarista, only A. paraguayensis also has similarly pubes- cent leaves. Agarista chapadensis is easily distinguished from A. subrotunda by its rounded- to cuneate- (vs. cordate-)based, often longer leaves and by its shorter pedicels; the taxa are geographically separated (Map 5). 1984] JUDD, AGARISTA 307 13. Agarista paraguayensis (Sleumer) Judd, comb. nov. Leucothoé paraguayensis Sleumer, Bot. Jahrb. 78: 465. 1959. Type: Paraguay, Para- guari, gees eae Piraveta, 23 Nov. 1950, Sparre & Vervoorst 501 (holotype, LIL; isotypes, C!, K!, s!). nie nas Sleumer var. ca/va Sleumer, Bot. Jahrb. 78: 465. 1959. Type: Paraguay, Amambay, Sierra de Amambay, Cerro Cora, Aug. 1933, Rojas 6189 (holotype, s1). Shrub to ca. 3 m tall. Twigs with or without scattered gland-headed hairs, otherwise sparsely to densely pubescent, with nonchambered to hollow or very sparsely septate pith. Buds to ca. 2.5 mm long. Leaves alternate; petiole 2.5- 6 mm long; blade revolute in bud, ovate, 1.8-5.2 by 0.7-2.2 cm, + flat, co- riaceous, the apex acute- to rounded-mucronate or slightly acuminate, the base rounded to cordate, the margin entire (slightly undulate), plane to very slightly revolute, especially near base, the adaxial surface pubescent on midvein (also with few unicellular hairs on lamina), the abaxial surface densely to irregularly and sparsely pubescent on midvein and lamina (glabrous or with pubescence limited to midvein), often with inconspicuous glandular dots along midvein. Inflorescences axillary racemes to 1.5-9 cm long, the axis with or without scattered gland-headed hairs, otherwise moderately to densely pubescent. Ped- icels 3-9 mm long, with or without gland-headed hairs, otherwise densely to moderately oe pubescent; bracteoles 2, opposite to alternate, from nearly basal to within lower 5 of pedicel, narrowly triangular, to ca. 1.5 mm long; bracts to 1.5 mm ae Calyx lobes triangular with acuminate apices, 0.9-2.5 by 0.9-1.8 mm, the abaxial surface sometimes with few gland-headed hairs, otherwise sparsely to densely pubescent; corolla cylindrical, 6.5—9 by 3-4 mm, red to white(?), abaxially slightly to very slightly pubescent (to glabrous’), filaments 4.5-5.5 mm long, anthers 1.1-2 mm long; ovary nearly glabrous to densely pubescent. Capsules subglobose, 3.5-4.5 by 5-6.5 mm, placentae + central; seeds 0.9-1.8 mm long. DISTRIBUTION AND ECOLOGY. Eastern Paraguay and adjacent Argentina (MAP 6). Rocky areas, steep slopes, ravine slopes along arroyos. Flowering August to December. REPRESENTATIVE SPECIMENS. Paraguay: Campt. Yagin, Chodat s.n., 1914 (a); ie Tobaty, Cerro Tobaty, Schinini 7920 (L). Argentina: Misiones, dept. San Ignacio yucuaré, Pefién Reina Victoria, Schinini 5499 (F, L, MO); Teyucuare, Medan et al. 108 (BAA). This poorly known species is the only taxon of Agarista known to occur in either Paraguay or Argentina. It is probably allied with 4. oleifolia and A. pulchella, from which it can be distinguished by its distinctive combination of often abaxially pubescent leaves and corollas and more or less hollow pith. Agarista paraguayensis is superficially similar to A. mexicana var. pinetorum, A. revoluta var. velutina, A. chapadensis, and A. subrotunda, all of which have abaxially pubescent leaves. However, it differs from them in placenta position, seed length, and pith structure. This taxon is variable in leaf indumentum. The leaves of many individuals 308 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Map 6. Distribution of Agarista oleifolia var. oleifolia (circles), A. oleifolia var. glabra (dots), A. paraguayensis (solid triangles), A. minensis (open squares), A. pulchella var. pulchella (solid squares), and 4. pulchella var. cordifolia Goa tanec). have a more or less moderate to dense layer of unicellular hairs on the abaxial surface, but those of some plants (e.g., Sparre & Vervoorst 501) are essentially glabrous beneath. The taxonomic significance of this variation will remain unknown until the taxon is better known and has been studied in the field. 14. Agarista oleifolia (Cham.) G. Don, Gen. Syst. 3: 838. 1834. Shrub (to small tree) to 3(-6) m tall, with thickly furrowed bark and often tortuous branches. Twigs with or without scattered gland-headed hairs, oth- erwise glabrous to sparsely pubescent, with nonchambered to clearly chambered pith. Buds to ca. 1 mm long. Leaves alternate (to nearly 3-whorled); petiole 2- 11.5 mm long; blade revolute in bud, (very narrowly) ovate to elliptic or oblong, (2—)2.5-10(—12.5) by 0.4—3.3 cm, + flat, nea the apex acute- to rounded- mucronate (slightly acuminate), the base cuneate to rounded or slightly cordate, the margin entire (slightly undulate), plane to very slightly revolute, especially near base, the adaxial surface nearly glabrous to sparsely pubescent on midvein, especially proximal portion, the abaxial surface glabrous to sparsely pubescent 1984] JUDD, AGARISTA 309 on midvein, usually with inconspicuous glandular dots along midvein. Inflo- rescences axillary racemes to (2.5—)3-13 cm long, the axis with or without scattered gland-headed hairs, otherwise glabrous to densely pubescent. Pedicels 2.5-15 mm long, with or without gland-headed hairs, otherwise glabrous to densely pubescent; bracteoles 2 (rarely 3), alternate to subopposite, from nearly basal to near midpoint of pedicel (rarely with 1 near apex), to ca. 1.5 mm long; bracts to 2 mm long. Calyx lobes triangular with acuminate apices, 1.4-2.7(-3) by 0.8-2.1 mm, the abaxial surface with or without gland-headed hairs, otherwise glabrous to moderately pubescent; corolla cylindrical, 7.5-16 by (2.5-)3-6.5 mm, red to white, abaxially glabrous; filaments 5.3-8.5 mm long, anthers 1-2.2 mm long; ovary glabrous to sparsely pubescent. Capsules subglo- bose to short-ovoid, 2.5-5.5 by 4-8 mm, placentae + central; seeds 0.5-1.3 mm long. DiIsTRIBUTION. Southeastern Brazil, from Bahia, Goids, and extreme south- eastern Mato Grosso south to Estado do Rio and Sao Paulo KEY TO THE VARIETIES OF AGARISTA OLEIFOLIA 1. Inflorescence axis sparsely to densely pubescent; corolla 7.5-12 mm long. ........ ds ead vitatihe sintloten Acaksiau aed Whee oe SA do Ba ee 14a. var. oleifolia. 1. Inflorescence axis glabrous or essentially so; corolla 10.5-16 mm long. ........... ee Coe te eee ey See ree G2 nk Cee eee ene 14b. var. glabra. 14a. Agarista oleifolia (Cham.) G. Don var. oleifolia FIGURE 3, b. Andromeda oleifolia Cham. Linnaea 8: 504. 1833. Leucothoé oleifolia (Cham.) DC. Prodr. 7: 605. 1839. Type: Brazil, Minas Gerais, Itambé, Se//ow s.n. (holotype, 8 eer fragments of holotype, F!, Ny; photos of holotype, F, GH!; isotypes, E!, Gl, kl, LI). Leucothoe ambigua Meissner var. tomentella Meissner in Martius, Fl. Brasil. 7: 156. 3(= L. ee var. ambigua; chosen as nominal variety of A. ambigua by ane 1959, since Meissner described six named varieties of an taxon at the same time). pres ambigua (Meissner) J . D. Hooker ex Niedenzu, Bot. Jahrb. 11: 236. 1889. Type: Brazil, Minas Gerais, Martius 537 (lectotype, ae holotype of L. ambigua var. oe fragment of lectotype, Fs ee G!, GH!, K!, L!, m!, Mo!, Ny!, w; photos of isotype, B (destroyed), G!, GH!). Leucothoé ambigua Meissner var. hispidula Meissner in ee s, Fl. Brasil. 7: 156. 1863. Leucothoé Cham.) DC. var. hispidula (Meisna) ee Bot. Jahrb. 78: 476. 1959. TYPE: Brazil, Minas Gerais, Carunhanha R., Sertao do Paranan, Martius 2000 (holotype, M!). Leucothoé ambigua Meissner var. peduncularis Meissner in Martius, Fl. Brasil. 7: 156. 1863. Type: Brazil, Minas Gerais, Serra do Frio, July 1846, Gardner ees, BM orcas fragment of holotype, Ny!; isotypes, E!, F!, G!, GH!, K!, Ny!, Pl, s us!, bevels stenophylla Loesener, Flora 72: 77. 1889. Agarista le as (Loesener) Niedenzu, Bot. Jahrb. 11: 236. 1889. Type: Brazil, Estado do Rio, Serra dos Orgaos 21 Jan. 1887, Glaziou 16232 (holotype, B (destroyed); Oe. . ‘st here designated as lectotype; isolectotypes, c!, F!, K!, P!). Leucothoé dang Sleumer, Notizbl. Bot. Gart. Berlin 12: 481. 1935. Type: Brazil, Minas Gerais, Serra do Itatiaia, 2100 m, 27 Dec. 1895, Ule 3737 (holotype, 8 (destroyed); eer of holotype, F!; photos of holotype, Fl, GH!; isotype, HBG!). 310 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Shrub (to small tree) to 3(-6) m tall. Twigs glabrous to sparsely pubescent. Leaves with petiole 2-8 mm long; blade (2-)2.5—8.5(-10.5) by 0.4-2.8 cm, base cuneate, abaxial surface glabrous to sparsely pubescent on midvein. In- florescences (2.5—)3-12 cm long, axis sparsely to densely pubescent. Pedicels glabrous to densely pubescent. Corolla 7.5—12 mm long, red to white; filaments 5.3-6.8 mm long, anthers 1-1.5 mm long; ovary glabrous to sparsely pubescent. Capsules 2.5—4 by 4-6.5 mm. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, southern and central Minas Gerais and adjacent portions of Estado do Rio and Sao Paulo (Map 6). Margins of forests and thickets, frequently along rivers or streams, bogs, open rocky campo; 800-2100 m alt. Flowering chiefly August through October ENTATIVE SPECIMENS. Brazil. Estapo Do Rio: Itatiaia, Brade 21265 (L); Novo Faburea, Capell s.n., 26 Nov. 1953 (L); Serra do Itatiaia, Retiro, Dusén 529 (s); Mont- serrat, Dusén 2124 (Gn, s); Parque Nacional do Itatiaia, margins Rio Campo Belo, Maas & Martinelli 3162 (Ny); Itatiaia, Abrigo Reboucas, Santos 5755 (L). Minas GERAIS: Andrelandia, Fazenda da Parahyba, Barreto 5276 (F, L, Ny); Rio Viravinha, Burchell 5495 (GH, K); Cachoeira do Campo, Casaretto 2861] (G); Curvello and Rio San Francisco, Claussen s.n., 1837 (L), Fazenda de Manoel José, Damazio 960 (c); Tiradentes, Duarte 835 (c, NY, 8, US); near Villa Rica, Itacolumi, Riedel 408 (G, GOET, K, L); Pico da ees near Caparaé, Shepherd et al. 5799 (F, L); Serra de Curimatai, St.-Hilaire, Cat. B', 1997 (p); Serra do Caparaé, Macieiras (Grotao), Strang 220 (1); Serra do Ouro Préto, Ule 2619 (HBG); aoe [Diamantina], Vauthier 5 (G, Gu, p). Sko PAULO: Moéca camp, Brade 5667 (s, sp), rd. from Sao Bernardo to Sao Paulo, Burchell 4054 (k); Butantan, Gehrt, SP n. 2083 (sp); Cidade Jardim, Kuh/mann, SP n. 35241 (sp); Campos do Jordao, Leite 3615 (A, Gu); Serra da Bocaina, S of Itatiaia, Markgraf & Aparicio 10418 (1) 14b. —— oleifolia (Cham.) G. Don var. glabra (Meissner) Judd, comb. FiGure_ 5, a, b. Leucothoé ambigua Meissner var. glabra Meissner in Martius, Fl. Brasil. 7: 156. 1863. Leucothoé oleifolia (Cham.) DC. var. glabra (Meissner) Sleumer, Bot. Jahrb. 78: 476. 1959. Type: Brazil, Bahia, ca. Moritiba, Serra da Jacobina, 1842-1845, Blanchet 3562 MoloW De, Ww; isotypes, BR, c!, F(specimen and photo)!, G!, GH(specimen and photo)!, k!, Mo!, p! Leucothoé ae Meissner var. longifolia Meissner in Martius, Fl. Brasil. 7: 156. 1863. Leucothoé oleifolia (Cham.) DC. var. /ongifolia (Meissner) Sleumer, Bot. Jahrb. 78: 476. 1959. Type: Brazil, Goiés, Arrayas, 1841, Gardner 3876 (holotype, BM; isotypes, E!, F!, G!, K!, Ny!, P!, w). Leucothoé martii Meissner var. puberula Meissner in Martius, Fl. Brasil. 7: 155. 1863. Brazil, Minas Gerais/Pernambuco, Rio Fermo, a aS: (holotype, M!; fragment of holotype, Ny; photo of holotype, Gu!; isotypes, G Minas Gerais/Pernambuco, Rio Fermo, Martius s.n. (holotype, m!; isotype, Leucothoé spectabilis Meissner in Martius, ibid. 159. Agarista spectabilis ae J. D. Hooker ex Niedenzu, Bot. Jahrb. 11: 236. 1889. Type: Brazil, Goids, banks of S. spe aaa Chapada de S. Marcos, Aug. 1834, Riede/ 2484 (holotype, LE; isotype, N 1984] JUDD, AGARISTA 311 d Ficure 5. a, b, Agarista el oe var. glabra (Irwin & ge acct rae in b note glabrous inflorescence axis an wers articulated with pedicels. c, pulchella var. nen (Hatschbach 19703): ae elongate racemes and oe a leaves. d, A minensis (Ule 1840): note short racemes and round-based leaves. Scale = 2c 312 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Leucothoé varnhageniana Reichardt, Verh. Zool.-Bot. Ges. Wien 33: 323. 1884. Aga- rista varnhageniana (Reichardt) Niedenzu, Bot. Jahrb. 11: 236. 1889. Type: Brazil, Goias, Villa Formosa do Imperatriz, Seguro 68 (holotype, w; fragment of holotype, F!; photos of holotype, a!, F!). Shrub (to small tree) to 3(-4) m tall. Twigs glabrous. Leaves with petiole 4— 11.5 mm long; blade (2.5—)3.5-10(-1 2.5) by 0.9-3.3 cm, base cuneate to round- ed, abaxial surface glabrous. Inflorescences 6-13 cm long, axis glabrous or essentially so. Pedicels glabrous. Corolla 10.5-16 mm long, red (rarely white); filaments 6.5-8.5 mm long, anthers 2-2.2 mm long; ovary glabrous. Capsules 3.5-5.5 by 5.5-8 mm DISTRIBUTION AND ECOLOGY. South-central Brazil, from Bahia and northern Minas Gerais east to Goids and southeastern Mato Grosso (Map 6). Scrub vegetation over sandstone, woodland and thickets, especially along rivers or streams; ca. 700-1700 m alt. Flowering chiefly August through October. REPRESENTATIVE SPECIMENS. Brazil. BAHIA: Serra do Rio de Contas, 3 km N of town of Rio de Contas, Harley et al. 15383 (£, L, NY, US); Serra do Sincora, by Rio Cumbuca, 3 km N of Mucugé on rd. to Andarai, Harley 18705 (xy); Serra do Tombador, 18 km E Irwin et al. 9852 (F, Ny); along Rio de Passa Quatro, Joaquim Diaz, St.-Hilaire, Cat. C, n. 869 (L, Pp). Mato Grosso: Rib. Claro, Alto Araguaia, Hatschbach 35079 (L). MINAS Gerais: near Grao Mogol, Williams & Assis 8190 (Gu). Agarista oleifolia is probably most closely related to 4. pulchella, A. deihetinns and A. paraguayensis. All four species have moderate to large, flat leaves an capsules with more or less centrally positioned placentae. Agarista oleifolia | 1s easily distinguished from A. paraguayensis by its abaxially glabrous corollas and leaves and its solid to chambered pith, and from 4. minensis by its longer inflorescences and often wider leaves. Agarista oleifolia is geographically sep- arated from both of these species. It can be distinguished from the very similar but geographically separated A. pulchella var. pulchella by its differently shaped leaves and its often longer petioles; in addition, the sympatric 4. pulchella var. cordifolia can be readily distinguished by its usually subapical placentae and its smaller leaves. Individuals of A. o/eifolia are also sometimes confused with A. uleana, A. niederleinii var. acutifolia, and A. coriifolia; see distinguishing characters in key. Populations of Agarista oleifolia are separable into two morphologically distinctive and geographically separate varieties. The more northern, var. g/a- bra, is best distinguished from var. oleifolia by its glabrous (vs. sparsely to densely covered with short unicellular hairs) inflorescence axis and its usually larger, more frequently red corollas. Variety o/eifolia has been illustrated by Meissner (1863, ¢. 58) Both taxa are variable in glandular indumentum and leaf shape, which has led to the description of numerous species and varieties (see synonymy) that cannot be maintained when the entire range of variation is considered. Indi- 1984] JUDD, AGARISTA 313 viduals with small, very narrowly ovate, acute-apexed leaves are frequent in the Serra do Itatiaia (and also occur in the Serra dos Orgaos). Such plants were described as Leucothoé stenophylla by Loesener (1889) and maintained as a distinct species by Sleumer (1959); however, this form intergrades completely with more typical A. oleifolia (see Dusén 2124 (GH, Ss), Dusén s.n., 15 June 1902 (s), Dusén s.n., 22 July 1902 (s)). Such narrow-leaved plants have oc- casionally been confused with A. minensis. Several specimens from Sao Paulo (i.e., Hoehne, SP no. (sp), Lutz 770 (ny, us), Lutz 1920 (L)) have small, ovate, + revolute, abaxially curved leaves, and Lutz 770 has robust capsules with + subapical placentae. These individuals may be hybrids between Agarista oleifolia and a species with small, strongly revolute leaves such as 4. hispidula or A. chlorantha. Leite 3615 (A, GH) has slightly ferrugineous hairs on its inflorescence axes and may represent a hybrid with the sympatric A. eucalyptoides. 15. Agarista minensis (Glaz. ex Sleumer) Judd, comb. nov. Ficure 5, d. Leucothoé minensis Glaz. Bull. Soc. Bot. France 57: 129. 1910, nomen nudum. Leu- cothoé minensis Glaz. ex Sleumer, Notizbl. Bot. Gart. Berlin 12: 480. 1935. Type: Brazil, Minas Gerais, Biribiry, Mocoté, near Diamantina, 28 March 1892, Glaziou 19572 (holotype, B (destroyed); isotype at K! here designated as lectotype; isolec- totype, P!). Shrub to 4 m tall. Twigs glabrous to sparsely pubescent, with clearly cham- bered pith. Buds to ca. 0.8 mm long. Leaves alternate; petiole 2-6 mm long: blade revolute in bud, slightly and narrowly ovate to narrowly oblong, (1-)1.6- 5.6 by 0.3-1(-1.7) cm, + flat, coriaceous, the apex acute- to rounded-mucro- nate, the base cuneate to truncate, the margin entire, plane to very slightly revolute, especially near base, the adaxial surface sparsely to very sparsely pubescent on proximal portion of midvein, the abaxial surface glabrous (with w very inconspicuous glandular dots along midvein). Inflorescences (fasci- clelike) axillary racemes to 0.5-1.5(-2.5) cm long, axis sparsely to densely pubescent. Pedicels 4-10 mm long, very sparsely to moderately pubescent: bracteoles 2, alternate to subopposite, from nearly basal to near midpoint of pedicel, narrowly triangular to linear, to ca. 1.6 mm long; bracts to 1.2 mm long. Calyx lobes triangular with acuminate (to acute) apices, 1.4-2.6 by 0.8- 1.4 mm, abaxial surface glabrous to sparsely pubescent; corolla cylindrical, 6.5-11 by 2.5-4.5 mm, white, abaxially glabrous; filaments 5-5.5 mm long. anthers ca. 1.2 mm long; ovary glabrous to very slightly pubescent near apex. Capsules short-ovoid to subglobose, 3-5 mm by 4.5-7.5 mm, placentae + central; seeds 1-1.5 mm long. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, Santa Catarina and Rio Grande do Sul (Map 6). Scrub and forests along streams and rivers; 900-1300 m alt. Flowering November through December (January). REPRESENTATIVE SPECIMENS. Brazil. Rio Sears bo SuL: Bom Jesus, Fazenda Bernardo Velho, Rambo 34939 (mo, s); Cambar tS. Francisco de Paula, Rambo 36728 (us): Taimbé, near S. Francisco de Paula, Ranibe 49306 (s); Jaquirana, near S. Francisco de 314 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Paula, Rambo 51933 (s, us); Passo do Inferno, near Canela, Rambo 56563 (B). SANTA CATARINA: Bom Jardim, Sao Joaquim, Reitz & Klein 7968 (L); Campo Alegre, Pinheiral, Morro Iquererim, Smith & Klein 7968 (L); 2 km W of Rio Capetinga on rd. to Dionisio Cerqueira, Smith et al. 9622 (us); Serra Geral, Campos de Capivari, Ule 1840 (upc, L). Although this taxon was included in Agarista niederleinii by Sleumer (1959), the two species differ greatly in capsule morphology and inflorescence length and are probably not closely related. Agarista minensis is actually closely related to A. pulchella var. pulchella, from which it is distinguished by its shorter inflorescences and its different leaf shape (length/width quotient (2.5-)3-6 vs. (1.4-)1.6-3(-3.5), base cuneate to truncate vs. cordate). (See Emrich & Rambo, 1949, fig. 20, and Marques & Klein, 1975, pi. 6, fig. 9.) The ranges of the two taxa overlap only slightly (Map 6). However, some hybridization may be oc- curring between them because a few specimens of A. pulchella (e.g., Smith & Klein 8470 (1, us), from Santa Catarina) show some A. minensis-like characters. Agarista minensis 1s also allied to A. oleifolia and can easily be confused with especially small and narrow-leaved individuals of 4. oleifolia var. oleifolia. The two can be distinguished by the length of the inflorescences and shape of the leaves—especially the apex, which is never narrowly acute in A. minensis but is usually so in narrow-leaved variants of A. oleifolia var. oleifolia. The provenance of the type (G/aziou 19572) is very uncertain. These spec- imens are identical with Ule 1840 (HBG, L) and 184] (see Sleumer, 1959), collected in the state of Santa Catarina. However, Glaziou 19572 was suppos- edly collected in Minas Gerais near Diamantina! Glaziou 19572 is the only collection of A. minensis from Minas Gerais, an area separate from the major portion of the species’ range, Santa Catarina and Rio Grande do Sul. It is very probable that the specimens represented by Glaziou 19572 were actually sent to Glaziou by Ule, and that the label information was altered by Glaziou; see discussion in Wurdack (1970). 16. Agarista pulchella Cham. ex G. Don, Gen. Hist. 3: 838. 1834. Shrub to 2(-3) m tall. Twigs with or without scattered gland-headed hairs, otherwise essentially glabrous to densely pubescent, with nonchambered to irregularly chambered pith. Buds to ca. | mm long. Leaves alternate; petiole 1.5-5 mm long; blade revolute in bud, + ovate, 0.5-4(-5) by 0.4—2(-2.8) cm, + flat, coriaceous, the apex rounded- to acute-mucronate (short-acuminate), the base cordate (to rarely truncate), the margin entire (to serrulate due to gland-headed hairs) (undulate), plane to very slightly revolute, especially near base, the adaxial surface glabrous to sparsely pubescent on midvein, especially proximal portion, the abaxial surface with or without few gland-headed hairs on midvein, otherwise glabrous to sparsely pubescent (with few very incon- spicuous glandular dots) on midvein. Inflorescences axillary racemes (axillary panicles, or terminal racemes or panicles) to (1.5—)2—7(-12) cm long, the axis with or without scattered gland-headed hairs, otherwise glabrous to densely pubescent. Pedicels 4-15 mm long, with or without gland-headed hairs, oth- erwise glabrous to densely pubescent; bracteoles 2 (to 5), alternate to subop- posite, from nearly basal to near midpoint (or rarely apex) of pedicel, narrowly 1984] JUDD, AGARISTA S15 triangular to linear, to ca. 2.2 mm long, occasionally with | or more subtending axillary flowers; bracts to 3 mm long. Calyx lobes triangular with acuminate (to acute) apices, 1.4-2.6 by 0.8-1.5 mm, the abaxial surface with or without gland-headed hairs, otherwise glabrous to densely pubescent; corolla cylindri- cal, 6.5-13 by 3-5.5 mm, white with reddish apex to red throughout, abaxially glabrous to sparsely pubescent; filaments 4-6.5 mm long, anthers 1.2-1.5 mm long; ovary glabrous to densely pubescent. Capsules short-ovoid to subglobose, 3-5 by 4.5-8 mm, placentae central to subapical; seeds 0.8-1.7 mm long. DISTRIBUTION. Southeastern Brazil, from Parana and Santa Catarina, and dis- junctly to the north in Minas Gerais and adjacent Sao Paulo. KEY TO THE VARIETIES OF AGARISTA PULCHELLA —_ Capsules with placentae usually + central; leaves with length/width quotient (1.4-)1.6-3(-3.5), seas a (rarely slightly) undulate; [Parana, Santa Catarina, and adjacent portion of Sao Tle nueugeee ooacte Aaa ace cue 6a. var. pulchella. . Capsules with placentae | ae subapical; leaves with length/width quotient 0.8-2, margin often clearly undulate; [Minas Gerais and adjacent northern portion of Saio WO) 5 cdl hea Miele uo eat nd Su 8 Maw Gini el os ea eae 16b. var. cordifolia. — 16a. Agarista pulchella Cham. ex G. Don var. pulchella Ficure_ 5, c. Andromeda pulchella Cham. Linnaea 8: 509. 1833, a later homonym of Andromeda pulchella Salisb. Prodr. 289. 1796, nomen superfl. (A. mariana L. = Lyonia mariana L.) D. Don cited in synonymy). Leucothoé pulchella (Cham. ex G. Don) DC. Prodi 7: 604. 1839. Type: Brazil, Minas Gerais, Antonio Pereira (see Meissner in Martius, 1863; locality very questionable since nearly all collections from Parana or Santa Catarina), Sellow 4830 (holotype, B (destroyed); fragments of holotype, F!, Ny; photos of holotype, F!, GH!). Shrub to 2(-3) m tall. Twigs glabrous to densely pubescent. Leaves with petiole 1.5-5 mm long; blade 1.2—4(—5) by 0.4—1.8(—2.8) cm, margin not (rarely slightly) undulate; adaxial and abaxial surfaces glabrous to sparsely pubescent on midvein. Inflorescences axillary racemes (axillary panicles, or terminal ra- cemes or panicles) to 2—7(—12) cm long, axis glabrous to densely pubescent. Corolla 7.5-13 by 3-5 mm; filaments 4.8-6.5 mm long. Capsules 3-4 by 5.5- 6 mm, placentae + central (to rarely subapical); seeds 0.8—1.3 mm long. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, Parana, Santa Catarina, and adjacent regions of Sao Paulo (Map 6). Gallery forests along rivers or streams, bogs, thickets or thicket-margins, sandy or rocky campo; (110-)780-1500 m alt. Flowering (August) September through November (December). EPRESENTATIVE SPECIMENS. Brazil. PARANA: Rio dos Papagaios, era 3009 & ae 2489 (L); Palmeira, Dombrowski 6596 (1); Serrinha, Dusén 6955 (F, GH, MO, s); Villa Velha, Dusén 7272 (s); Jaguariaiva, Jénsson 372 (A, s); Lapa, Rio Sao Vicente, Serrinha, Hatschbach (Curial) 518 (1, s); Lapa, Rio Passa Dois, Hatschbach 5097 e: Arapoti, Rio das Cinzas, Barra do Perdizes, Hatschbach 7223 (L); Palmeira, Rod. Café, acs 10158 (B); Pérto Amazonas, Fazenda Sao Luis, Hatschbach 10243 (). Ponta Grossa, Passo do Pupo, Hatschbach 17380 (L, uc); Balsa Nova, Barra Rio Pa- pagaios, erate 19703 (mo, uc); Estr. do Cerne, Serra das Furnas, Pirai do Sul, 316 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Hatschbach & Guimaraes 24761 (kK, MO, S, UC, US); Morro da Baliza, Lageadinho, Palmas, Hatschbach 30747 (c, L, uc, us); Serra da Canha, Cerro Azul, Hatschbach 32608 (c, L, uc); Rod. dos Mineiros, Almirante Tamandaré, Hatschbach 37224 (L); Estr. Curitiba to SAo PauLo: Itararé, Campos de Sao Pedro, near Serra de Bom Sucesso, Fazenda Ventania, Mattos 14108 (sp) 16b. Agarista pulchella Cham. ex G. Don var. cordifolia (Meissner) Judd, comb. nov. Leucothoé cordifolia Meissner in Martius, Fl. Brasil. 7: 162. t. 60, fig. 2. 1863. Agarista cordifolia (Meissner) J. D. Hooker ex Niedenzu, Bot. Jahrb. 11: 236. 1889. Leucothoé pulchella (Cham. ex G. Don) DC. var. se he (Meissner) Sleumer, Bot. Jahrb. 78: 473. 1959. Type: Brazil, Minas Gerais, Serra do Sao Joao d’El Rey and Serra de Sao José, June 1824, Riedel 308 (holotype, LE; isotype, NY!). Shrub to 1.5 m tall. Twigs moderately to densely pubescent. Leaves with petiole 1.5-3 mm long; blade 0.5—3.5 by 0.4—2 cm, margin often clearly un- dulate; adaxial and abaxial surfaces + sparsely pubescent on midvein. Inflo- rescences axillary racemes to (1.5-)2-5.5 cm long, axis densely pubescent. Corolla 6.5-10 by 3.5-5.5 mm; filaments 4-4.5(?) mm long. Capsules 2.5—5 by 4.5-8 mm, placentae usually subapical; seeds ca. 0.8-1.7 mm long. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, Minas Gerais and adjacent Sao Paulo (Map 6). Dry, rocky campo; ca. 1600-1700(?) m alt. Flowering March, June, July, and October through December. REPRESENTATIVE SPECIMENS. Brazil. MiNAs GERAIS: Serra de Tiradentes, Barreto 17476 (Ny); So José d’El Rey and Ouro Préto, Glaziou 17109 (B, c, Ny); Serra do Ibitipoca, St.-Hilaire, Cat. D, n. 238 (p). SAo Pauto: Serra da Bocaina, Brade & Duarte 21179 (L); Serra da Bocaina, Vacca Cahio, Glaziou 8236 (c, G, P, US); Barreiro Co., Serra da Bocaina, Lageado Farm, Segadas-Vianna 2800 (L); Silveiras Co., Serra da Bocaina, Jardim Farm, Segadas- Vianna 3256 (1). Agarista pulchella is most closely related to A. minensis and A. oleifolia. It can be distinguished from the former by its longer inflorescences and by the length/width quotient (usually 1.6-3 vs. 3-6) and the bases (cordate vs. cuneate to truncate) of its leaves; from the latter, by its often shorter, more consistently ovate, cordate-based leaves with often shorter petioles. The ranges of A. pul- chella and A. minensis overlap slightly (in northern Santa Catarina), and a few intermediate specimens are known (e.g., Smith & Klein 8470, L, us); thus, some hybridization may be occurring. Agarista oleifolia is geographically separated from the southern A. pulchella var. pulchella, but it occurs sympatrically with the northern var. cordifolia. Agarista pulchella var. cordifolia 1s easily separable from A. oleifolia by the characters given above, as well as by its subapical placentae. Intermediate specimens are apparently unknown. Specimens of 4. pulchella are occasionally identified as A. niederleinii, A. subcordata, or A. coriifolia; see distinguishing characters given in key. 1984] JUDD, AGARISTA 3] Populations of Agarista pulchella are divisible into two geographically sep- arate varieties (see illustration of var. pulchella in Marques & Klein (1975, pi. 7), and of var. cordifolia in Meissner (1863, t. 60, fig. 2)). Variety pulchella has been widely collected in Parana and Santa Catarina, while var. cordifolia seems to be uncommon and limited to a small portion of Minas Gerais and adjacent Sao Paulo (Map 6). Sleumer (1959) first recognized the close relationship be- tween these two taxa. Although the capsules of 4. pulchella var. pulchella usually have + central placentae, plants with subapical placentae (e.g., Jonsson 224, s)—the typical condition in var. cordifolia—are known. St.-Hilaire, Cat. D, n. 238 is here placed in A. pulchella var. cordifolia; this specimen is aberrant in having moderately pubescent corollas and capsules with central placentae. However, A. pulchella var. cordifolia sometimes has slightly pubescent corollas (see Riedel 308, Ny), and individuals with atypical placenta position occur very occasionally in the southern populations of this species. The unusual placenta position could possiply indicate Pat this specimen is a hybrid with A. hispidula, a species with central p t ly pubescent corollas that grows sympatrically with 4, pulchella in southern Minas Gerais. Field studies are necessary to determine the placement of this unusual specimen conclusively. Agarista pulchella var. pulchella is quite variable in the shape and the length/ width quotient of its leaves. At one extreme are plants with short, rather broad, clearly cordate-based leaves, and at the other are those with longer, narrower, and only slightly cordate ones. The former plants are vegetatively rather similar to var. cordifolia and have often been identified as such, while the latter are easily confused with A. minensis. 17. Agarista nummularia (Cham. & Schldl.) G. Don, Gen. Syst. 3: 837. FiGuRE 6, a, b. Andromeda nummularia Cham. & Schldl. Linnaea 1: 520. 1826. Leucothoé num- mularia (Cham. & Schldl.) DC. Prodr. 7: 603. 1839. Type: Brazil, Rio Grande do Sul, Pérto Alegre (see Meissner in Martius, 1863), Sellow 1229 (holotype, B (de- stroyed); fragment of holotype, F!; photos of holotype, F!, G!, GH!; isotype, G-pc). Leucothoé nummularia (Cham. & Schldl.) DC. var. floccigera Sleumer, Bot. Jahrb. 78: 460. 1959. Type: Brazil, Rio Grande do Sul, Povo Novo, near Pelotas, 12 Nov. 1901, Malme 401 (holotype, s!; isotypes, Gul, s!). Erect shrub or subshrub to 2 m tall, with bark usually not well developed. Twigs with or without scattered gland-headed hairs, otherwise sparsely to densely pubescent, with hollow to irregularly chambered pith. Buds to ca. 1.2 mm long. Leaves alternate; petiole 0.5—2.5 mm long; blade revolute in bud, ovate to elliptic or orbicular, 0.5-1.5(—2.3) by 0.5—1.6(-2) cm, + flat, coriaceous, the apex obtuse- to retuse-mucronate (rarely short-acuminate), the base cordate to + truncate (rounded), the margin entire to serrulate due to gland-headed hairs (slightly undulate), plane to very slightly revolute near base, the adaxial surface with or without gland-headed hairs, otherwise sparsely pubescent on proximal portion of midvein, the abaxial surface with or without gland-headed hairs on midvein and blade, otherwise glabrous to sparsely pubescent on proximal por- tion of midvein (with few very inconspicuous glandular dots along midvein). 318 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 ya biel ARS (Chipeta /182Z rium Anchieta Bastion australis il TRIC 1} 2 TMNTTTTTTNTTTTTI en d FiGure6. a,b, Agarista nummularia: a, Rambo 53881,b, Rambo 48695. c, A. virgata (Duarte 8098): note wandlike, rigidly ascending branches. d, 4. chlorantha (Ratter et al. 3384). Scale = 2 cm JAROIM BOT 1984] JUDD, AGARISTA 319 Inflorescences axillary racemes to 1.5-8 cm long, the axis with or without scattered gland-headed hairs, otherwise densely pubescent. Pedicels 4-11 mm long, with or without gland-headed hairs, otherwise densely pubescent, brac- teoles 2, alternate to opposite, from nearly basal to within lower ' of pedicel. narrowly triangular to linear, ovate, or elliptic, to ca. 2.5 mm long; bracts to 3 mm long. Calyx lobes triangular with acuminate to acute apices, (2.1—)2.5— 5 by 0.7-1.7(-2.2) cm, the abaxial surface with or without gland-headed hairs, otherwise sparsely to densely pubescent; corolla cylindrical, 7-10.5 by 2.5-5 mm, white, abaxially glabrous; filaments 4.5-5.5 mm long, anthers |-1.7 mm long; ovary densely pubescent. Capsules short-ovoid to subglobose, 3.5—5 by 5-7 mm, placentae subapical; seeds 1.2-2.3 mm long. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, Santa Catarina and Rio Grande do Sul (Map 7). Rocky areas, bogs, wet forests, and moist thickets; 10- 1650 m alt. Flowering (August) September through December (January). REPRESENTATIVE SPECIMENS. Brazil. Rio GRANDE DO SuL: Montenegro, Camargo 1786 (s); Morro Sapucaia, S40 Leopoldo, Leite 3257 (Gu); Cachoeira, Malme 401/a (s); Apa- rados da Serra, Pabst 6301 & Pereira 6474 (L), Cambar4, S. Francisco de Paula, Rambo 36724 (B, MO, NY, US); Gravatai, summit Monte Sapucaia, Rambo 42757 (B, L, Us); Sapucaia, S40 Leopoldo, Rambo 48695 (B, us); S. Francisco de Paula, Rambo 52951 (s); Serra da Roginha, near Bom Jesus, Rambo 53881 (s); Fazenda Englert, near S. Francisco de Paula, Rambo 54674 (s); Canela, Richter 7837 (L). SANTA CATARINA: Serra do Oratorio, Bom Jardim, S40 Joaquim, Reitz & Klein 7456 (L); Bom Retiro, Fazenda Santo Antonio, Campo dos Padres, Smith & Reitz 10333 (us), Sombrio, Reitz 1926 (G, S, US). Agarista nummularia is probably closely related to A. chlorantha and A. virgata; it can be separated from the former by its more or less flat, nonrevolute leaves, and from the latter by its sparsely to densely pubescent inflorescence axis, pedicels, and ovaries, and its capsules with more or less subapical pla- centae. These species are essentially geographically isolated. Agarista num- mularia may also be confused with small-leaved individuals of A. pulchella, a species with shorter calyx lobes that grows to the north of the range of A. nummularia (Maps 6, 7). The species has been illustrated by Marques and Klein (1975, pi. 10). 18. Agarista virgata Judd, sp. nov. FIGuRE 6, c. Frutex erectus ad ca. | m altus. Ramuli hornotini pilis glandulosis praediti, aliter glabri, cum medulla non septata. Folia ovata vel elliptica, 0.7—1.8 cm longa, 0.7-1.7 cm lata, coriacea, ad apicem brevissime acuminata, acuta vel obtusa cum mucrone brevi, ad basin rotundata vel cordata; margo planiuscula, integra vel minute serrulata; pagina abaxialis pilis glandulosis praedita (prae- cipue in nervo primario), aliter glabra. Inflorescentiae axillares vel terminales, racemosae vel paniculatae, ad 5—16 cm longae; axis pilis glandulosis praeditus, aliter glaber. Pedicelli 2-7 mm longi, pilis glandulosis praediti, aliter glabri. ores 5-merus. Calyx lobis 1.4-3 mm longis, 0.6-1.6 mm latis, cum pilis glandulosis in pagina abaxiali. Corolla cylindrica, 6-7 mm longa, 2-3 mm lata, alba, glabra in pagina abaxiali. Filamenta ca. 4 mm longa; antherae ca. | mm 320 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Distribution of Agarista nummularia (open squares), A. virgata (solid tri- angle), nm chlorantha (dots), A. organensis (solid square), A. hispidula (circles), and A. ericoides (open triangles). longae. Ovarium glabrum. Capsula subglobosa vel brevissime ovoidea, 4—5.5 mm longa, 5.5—7.5 mm lata, cum placentis + centralibus. Semina 1.2—1.6 mm longa. Erectly branched shrub to ca. | m tall. Twigs with scattered gland-headed hairs, otherwise glabrous, with nonchambered pith. Buds to ca. 2.8 mm long. Leaves alternate; petiole 1-2 mm long; blade revolute in bud, ovate to elliptic, 0.7-1.8 by 0.4-1.3 cm, + flat, coriaceous, the apex acute- to obtuse-mucronate to short-acuminate, the base cordate to rounded, the margin entire or serrulate due to gland-headed hairs, plane, the adaxial surface with gland-headed hairs, otherwise very sparsely pubescent on extreme proximal portion of midvein, the abaxial surface with gland-headed hairs especially on midvein, otherwise glabrous, usually with few very inconspicuous glandular dots along midvein. Inflorescences axillary racemes or terminal racemes or panicles, to 5-16 cm long, the axis with scattered gland-headed hairs, otherwise glabrous. Pedicels 2-7 mm long, with gland-headed hairs, otherwise glabrous; bracteoles 2, al- ternate to opposite, from near apex to midpoint of pedicel, narrowly triangular 1984] JUDD, AGARISTA 321 to ovate, to ca. 2.3 mm long; bracts to 5 mm long (grading into leaves). Calyx lobes triangular with acuminate apices, 1.4-3 by 0.6-1.6 mm, the abaxial surface with gland-headed hairs, otherwise glabrous; corolla cylindrical, 6-7 by 2-3 mm, white, abaxially glabrous; filaments ca. 4 mm long, anthers ca. | mm long; ovary glabrous. Capsules subglobose to short-ovoid, 4—5.5 by 5.5— 7.5 mm, placentae + central; seeds |1.2-1.6 mm long. Type. Brazil, Minas Gerais, Serra do Cipé, km 140, 22 June 1964, A. P. Duarie 8098 (holotype, F!; isotypes, L!, s!). DISTRIBUTION AND ECOLOGY. Southeastern Brazil, Minas Gerais, Serra do Cipé (Map 7); ca. 1400 m alt. Flowering in June. REPRESENTATIVE SPECIMEN. Brazil. MINAS GERAIS: Serra do Cipé, km 131, Duarte 2682 (G, L) This distinctive species is easily recognized by its rigidly ascending, wandlike branches (from which the specific epithet is derived) with small, more or less flat, densely overlapping leaves. It is most easily confused with the geograph- ically separated Agarista nummularia (see Mar 7). However, A. virgata is easily separated from A. nummularia by its twigs, inflorescence axis, pedicels, and ovaries that completely lack unicellular hairs, and by its capsules with more or less centrally located placentae. This species is also allied with A. hispidula and A. chlorantha. It is of interest that both known collections of this taxon have gland-headed hairs; further collecting may reveal plants lacking these hairs, since their pres- ence is variable in all other species in which they occur. 19. Agarista chlorantha (Cham.) G. Don, Gen. Syst. 3: 838. 1934. FiGure 6, d. Andromeda chlorantha Cham. Linnaea 8: 508. 1833. Leucothoé chlorantha (Cham.) DC. Prodr. 7: 604. 1839. Type: Brazil, without definite locality, Se/low s.n. (holotype, Boia): fragment of holotype, F!; photos of holotype, F!, G!, GH!; fragment of isotype, NY! Amechania subcanescens DC. Prodr. 7: 579. 1839. Leucothoé subcanescens (DC.) Meissner in Martius, Fl. Brasil. 7: 163. ¢. 62, fig. 1. 1863. Leucothoé chlorantha (Cham.) DC. var. subcanescens (DC.) cena Bot. Jahrb. 78: 454. 1959. Type: Brazil, Sdo Paulo, Batatais, June 1834, Lund s.n. (holotype, G-pc; fragment of holotype, NY; isotype, Cc Andromeda serrulata Cham. Linnaea 8: 506. 1833. Agarista serrulata (Cham.) G. Don, Brazil, Sao Paulo (see Meissner in Martius, 1863), Sel/low s.n. (holotype, B (de- stroyed); fragments of holotype, F!, Ny; photo of holotype, Gu!; isotypes, E!, G!, K!). Erectly branched shrub or subshrub to 1.5 m tall, with bark usually not well developed. Twigs with or without scattered gland-headed hairs, otherwise mod- erately to densely pubescent, with + nonchambered to hollow pith. Buds to ca. 0.6 mm long. Leaves alternate; petiole 1-4 mm long; blade revolute in bud, ovate or narrowly ovate to orbicular or even suborbicular, 0.7-2.5 by 0.25- 1.3(-1.7) cm, strongly to slightly abaxially curved (rarely + flat), coriaceous, Se JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 the apex acute- to rounded-mucronate, the base cordate, the margin entire, often undulate (serrulate due to gland-headed hairs), strongly to slightly revolute (rarely + plane), the adaxial surface sparsely pubescent on midvein, especially proximal portion, the abaxial surface with or without gland-headed hairs on midvein, otherwise glabrous to sparsely pubescent on proximal portion of midvein, often with few very inconspicuous glandular dots along midvein. Inflorescences axillary (rarely terminal) racemes to (1—)1.5—8.5 cm long (very rarely flowers solitary, axillary), the axis with or without scattered gland-headed hairs, otherwise moderately to densely pubescent. Pedicels 3.5-13 mm long, with or without gland-headed hairs, otherwise moderately to densely pubescent; bracteoles 2 (rarely several), alternate to subopposite, from nearly basal to within lower 3 of pedicel (rarely to near apex), triangular to linear (ovate), to ca. 2.5 mm long; bracts to 3 mm long. Calyx lobes triangular to ovate, with + acute apices, 2.5—5.5 by 0.8-1.7 mm, the abaxial surface with or without gland- headed hairs, otherwise very sparsely to densely pubescent; corolla cylindrical, 7-11.5 by 3-6.5 mm, white (reddish toward apex), abaxially glabrous; filaments 5.5-6.5 mm long, anthers 1—1.6 mm long; ovary sparsely to densely pubescent. Capsules short-ovoid to subglobose, 3-5 by 4.5-7 mm, placentae subapical; seeds 1—2.1 mm long DISTRIBUTION AND ECOLOGY. Southeastern Brazil, from Distrito Federal south to Minas Gerais and Sao Paulo, and disjunctly from Parana south to Santa Catarina (Map 7). Wet, boggy or marshy campo, with grasses, sedges, and Xyris, open rocky and/or grassy areas, gallery forest; 700-1650 m alt. Flowering late August through December (January). REPRESENTATIVE SPECIMENS, Brazil. Dist. FEDERAL: Cérrego Samambaia, near Taguatin- ga, Irwin et al. 8151 (Ny, sp, UC, US); Rio Torto, N of Brasilia, Irwin et al. 8433 (Ny), 3050 (cE). PARANA: Campos do Capao da Imbuia, Curitiba, Dombrowski 408 & Saito 227 (A, L); Col6nia Orledes, Curitiba, ne 3088 (L); Pinhaes, Dusén 7082 (GH, MO, NY, S); Piraquara, Rio Palmital, Hatschbach 781 (.); Palmas, Sta. Barbara, Hatsch- bach 15005 (F, L, NY, UC, US, wis); Rio Pequeno, S. José dos Pinhais, Hatschbach 22815 (c, L, Uc); S. Joao do Triunfo, Hatschbach 17748 (L, uc); Clevelandia, Hatschbach 22714 (L, uc); S. Jer6nimo da Serra, Rio S. Jerénimo, Hatschbach & Guimardes 24777 (c, s uc); Rio Atuba, Curitiba, Hatschbach 32738 (L); Rio Palmital, Colombo, Halschbach 32792 (L, MO, Uc); Uniao da Vitéria, Koczicki 48 (L, uc); Tatuquara, Curitiba, Kummrow 692 (L). MINAS GERAIS: Uberaba, Lund s.n., 1843 (c). SANTA CATARINA: Fazenda Car- neiros, Cacador, Klein 3546 (1); Planalto Catarinense, Pereira 6290 & Pabst 6117 (1); Morro do Iquererim, Campo Alegre, Reitz & Klein 5220 (L, us); Serra da Boa Vista, Sao José, Reitz & Klein 5418 (L, us); Ponte Alta do Norte, Curitibanos, Reitz & Klein 13386 (L); Valdes, Reitz & Klein 13549 (L); Fazenda Frei Rogério, Pérto Unido, Reitz & Klein 13609 (L); Campo do Areao, Santa Cecilia, Reitz & Klein 14196 (L); Fazenda Ernesto Scheide, Campo Alegre, Reitz & Klein 5324 (L, us); Bom Retiro, falls of Rio Canoas, Campo dos Padres, Smith & Klein 7861 (us); Chapec6, Fazenda Campo Sao Vicente, 24 km W of Campo Eré, Smith et al. 9525 (L, us); Bom Retiro, Fazenda Santo Ant6nio, Campo dos Padres, Smith & Reitz 10331 (L, us); Irani, Campo de Irani, Smith & Klein 13032 (us); Agua Doce, Campos de Palmas, 3 km NW of Hercilidpolis, Smith & Klein 13634 (GH, L, NY, UC, US); 6 km W of Campo Eré, Smith & Klein 13708 (L, Ny, US). SAO PauLo: Campos do Jordao, Hashimoto 286 (sp), Leite 3942 (A, GH); Butantan, Hoehne, 1984] JUDD, AGARISTA 320 SP n. 467 (sp), 572 (sp); SalesOpolis, near Rio Coruja, Mattos, SP n. 157944 (ny): Araramara, St.-Hilaire, Cat. C! n. 1004 (rp); Urupunga, St.-Hilaire, Cat. C' n. 1048 (P). Agarista chlorantha is most similar to (and easily confused with) A. hispidula and A. organensis; all three taxa are characterized by more or less erect branches with small, usually ovate, clearly revolute leaves. This species is easily separated from A. hispidula by its abaxially glabrous (vs. pubescent) corollas and its more or less acute (vs. acuminate) calyx lobes. It can be distinguished from 4. or- ganensis by its longer calyx lobes, corollas, and inflorescences, although a few specimens (e.g., Dombrowski 408, Leite 3942, Mattos, SP n. 157944, Reitz & Klein 5220) have very short inflorescences or rarely even solitary flowers and thus somewhat approach the latter species. Agarista chlorantha is geographi- cally separated from A. organensis (see Map 7), although it grows together with A. hispidula in the northern portion of its range (Map 7). Specimens inter- mediate between A. chlorantha and A. hispidula are apparently unknown, but field studies in their area of overlap would be of interest. Agarista chlorantha appears to be more common (or at least much better collected) in the southern portion of its range (Parana and Santa Catarina). Agarista chlorantha occurs sympatrically with 4. pulchella over much of its range, and Hatschbach 17748 (L, uc) may represent a hybrid between these species. This specimen is unusual in its large (to 3.7 by 2 cm) nonrevolute leaves Like many other Agarista species, 4. chlorantha is variable in its glandular indumentum. There may be many to few multicellular, gland-headed hairs on the twigs, leaves, inflorescence axis, pedicels, and calyx lobes, or these hairs may be lacking. The plants are alike in all other features, and both indument forms may be found in a single locality (e.g., Smith & Klein 13634, Hoehne, SP n. 467, 572). Thus, A. serrulata and A. chlorantha, the former characterized by Sleumer (1959) as being glandular-hairy and the latter as lacking such hairs, are considered to be conspecific. Leaves of Agarista chlorantha are also some- what variable in extent of marginal revolution: those near the base of the plant are frequently less strongly revolute than those toward the distal part of the shoots. In a few plants many of the leaves are only very slightly revolute and more or less orbicular; such plants are easily confused with the closely related A. nummularia, a species with consistently plane and widely ovate to elliptic or nearly orbicular leaves. The species has been illustrated several times—see Meissner (1863, ¢. 1, fig. 1. t. 62, fig. 1; t. 63, fig. 1) and Marques and Klein (1975, pl. 9). 20. Agarista organensis (Gardner) J. D. Hooker ex Niedenzu, Bot. Jahrb. 11: 236. 1889. FIGURE 7, C. Leucothoé organensis Gardner in W. J. Hooker, London Jour. Bot. 4: 132. 1845. Tyee: Brazil, Estado do Rio, Serra dos Orgaos, ca. 1675 m alt., May 1837, Gardner 475 (lectotype, K!; isolectotypes, F(fragment)!, G, Gu!, Ny!, P!; photos of isolectotype, F'!, G, GH!). Erect shrub to ca. 2 m tall. Twigs very sparsely to moderately pubescent, with nonchambered to clearly chambered pith. Buds to ca. 0.7 mm long. Leaves 324 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 FiGure 7. a, b, Agarista hispidula: a, Leite 3477. b, Dusen s.n., 22 ack 1902 (s). c, A, organensis (Glaziou 17119). d, A. ericoides (Glaziou 19577), Scale = 2 ¢ 1984] JUDD, AGARISTA 32) alternate; petiole 1.5-4 mm long; blade revolute in bud, ovate (to elliptic), 0.5- 1.7 by 0.15-0.8 cm, strongly abaxially curved, coriaceous, the apex acute- to rounded-mucronate, the base cordate to truncate, the margin entire, strongly revolute, the adaxial surface glabrous to sparsely pubescent on midvein, es- pecially proximal portion, the abaxial surface glabrous to sparsely pubescent on midvein, usually with few very inconspicuous glandular dots along midvein. Inflorescences axillary, fasciclelike, often only few flowered racemes to 0.3-1.5 cm long, or flowers solitary and axillary, axis moderately to densely pubescent. Pedicels 4-8 mm long, sparsely to densely pubescent; bracteoles 2 to several, alternate to subopposite, from near base to near midpoint of pedicel, narrowly triangular, to ca. 0.9 mm long; bracts to ca. 1 mm long. Calyx lobes triangular with acuminate apices, 1.1-2 by 0.5-1.2 mm, abaxial surface sparsely to mod- erately pubescent; corolla cylindrical, 6-7.5 by 2-4 mm, white to red(?), abax- ially glabrous; filaments 4—4.5 mm long, anthers 1—-1.3 mm long; ovary sparsely to moderately pubescent. Capsules short-ovoid, 3-5 by 4.5-6 mm, placentae subapical; seeds 1-3 mm long. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, Estado do Rio, Serra dos Orgios (Map 7). Shrub bogs with Sphagnum; ca. 1500-1830 m alt. Flowering November through January. REPRESENTATIVE SPECIMENS. Brazil. Estapo po Rio: Serra dos Orgaos, Gardner 5802 (k, Ny), Glaziou 3763 (c, K, L), Glaziou 16231 (A, B, C, F, NY, P), Glaziou 17119 (A, C, F, K, NY, UC, US), Guillemin 948 (P). Agarista organensis is very closely related to (and easily confused with) A. chlorantha. Both species are shrubs with more or less rigidly ascending branches bearing small, more or less ovate, usually moderately to strongly revolute, cordate-based leaves. Agarista organensis can be distinguished from A. chlor- antha by its short calyx lobes and its very short fasciclelike racemes or its solitary flowers. However, some fasciculate or solitary-flowered variants of A. chlorantha are known, and additional collections are necessary in order to clarify the taxonomic value of this character. Agarista organensis is geograph- ically isolated; within the genus only A. oleifolia also occurs in the Serra dos rgaos. 21. Agarista hispidula (DC.) J. D. Hooker ex Niedenzu, Bot. Jahrb. 11: 236. 1889. FiGurRE 7, a, b. Te hispidula DC. Prodr. 7: 579. 1839. Leucothoé hispidula (DC.) Meissner in s, Fl. Brasil. 7: 164. ¢. 62, fig. 2. 1863. Type: Brazil, Sao Paulo, eae June 1834, ats n. (holotype, G-pc; fragment of holotype, Ny; isotype, Leucothoé breviflora Meissner in Martius, ibid. 165. t. 63, fig. 2. ae breviflora (Meissner) J. D. Hooker ex Niedenzu, Bot. Jahrb. 11: 236. 1889. Type: Brazil, Minas Gerais, Caldas, Oct. 1854, Lindberg 418 (holotype, BR; fragment of holotype, Ny!; isotype, s!). Leucothoé intermedia Meissner in Martius, Fl. Brasil. 7: 163. t. 60, fig. 1. 1863. Type: Brazil, Minas Gerais, Caldas, 1845, Widgren 329 (lectotype (here designated), BR; fragment of lectotype, Ny!; possible isolectotype, s!). 326 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Usually erect shrub to 2.5 m tall (but often only ca. 1 m tall), with bark often not well developed. Twigs with or without scattered gland-headed hairs, oth- erwise sparsely to densely pubescent, with nonchambered to irregularly cham- bered or sometimes hollow pith. Buds to ca. 0.8 mm long. Leaves alternate; petiole 1-4 mm long; blade revolute in bud, ovate to narrowly ovate, 0.8-2.5(-3.3) by 0.15-1.2 cm, strongly to slightly abaxially curved, coriaceous, the apex acute- to rounded-mucronate (short-acuminate), the base cordate, the margin entire (slightly undulate) (serrulate due to gland-headed hairs), strongly to moderately (only slightly) revolute, the adaxial surface sparsely to densely pubescent on midvein and often with scattered hairs on lamina, the abaxial surface with or without gland-headed hairs on midvein, otherwise sparsely to densely pubescent on midvein and often also sparsely pubescent on lamina (with few very inconspicuous glandular dots along midvein). Inflorescences (very short, fasciclelike) axillary racemes (rarely terminal racemes or panicles) to 0.5-6.5 cm long, axis with or without scattered gland-headed hairs, otherwise densely pubescent. Pedicels S—13 mm long, with or without gland-headed hairs, otherwise moderately to densely pubescent; bracteoles 2 (rarely 3), alternate to eee ihe nearly basal to near midpoint of pedicel, narrowly triangular, ca. 1.8 mm long; bracts to 2.3 mm long. Calyx lobes triangular to ovate a ae (to rarely + acute) apices, 2-5.5 by 0.8-—2 mm, abaxial surface with or without gland-headed hairs, otherwise sparsely to densely pubescent; corolla cylindrical to urceolate-cylindrical, 6.5-8.5 by 3-6 mm, red to pink (or white), abaxially sparsely to densely pubescent (rarely also with few gland- headed hairs); filaments 4-5 mm long, anthers 1.4—1.8 mm long; ovary densely pubescent. Capsules short-ovoid, 2.5—5 by 4-7.5 mm, placentae subapical to central; seeds 0.6—1.4 mm long. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, southern Minas Gerais and adjacent Sao Paulo and Estado do Rio, with one specimen from Goids (MAP 7). Elfin woodlands, thickets, thicket margins, shrubby grassland, rocky sites, open boggy areas; 1000-2500 m alt. Flowering (June) September through De- cember (February). R Brazil. E Rio: Retiro, Dusén 5] (s); Itatiaia, Glaziou &78&7 (c, L, P); Serra do Itatiaia, W of Retiro de Ramos, aeons oes (s); summit, Mt. Itatiaia, Estagaéo Biologica, L. B. Smith 1504 (Gu, s); Planalto of Itatiaia, 150 km WNW of Rio de Janeiro, R. & A. Tryon 6696 (Ny); Serro do Itatiaia, Ule 3410 (upc); Parque Nacional de Itatiaia, Veloso 304 (Ny). GorAs: Corumba, Glaziou 21621 (P). MINAS GerAIs: Fazenda da Parahyba, Andrelandia, Barreto 5304 (L); Sao Thomé das Letras, Baipendi, Brade 20428 (F, M, Us); Pocos de Caldas, Emmerich 2140 (L); Camanducaia, Vila Monte Verde, Filho 1869 (L); Caldas, Cervo, Regnell III-838 (s); Caldas, between Rio Pardo and Corcovado, Regnell III-839 (p, s), Plateleiras, Agulhas Negras, Sucre 5767 (F). SAO PAULO: Campos do Jordao, Reserva do Inst. Forestal, Sdo José dos Alpes, Davis 3036 (£); S40 Carlos, Handro 401 (us); Campos do Jordéo, Kuhimann 32477 (sp), Leite 3477 (Aa, GH, 8); Moji-Guacu, Fazenda Campininha, 3 km NNW of Padua Sales, Mattos & Mattos 8214 (k). Agarista hispidula is most closely related to A. chlorantha, A. organensis, and A. ericoides; it can be distinguished from all of these species by its abaxially pubescent corollas. In addition, it differs from A. organensis in its elongated 1984] JUDD, AGARISTA 327 calyx lobes, from A. ch/orantha in its acuminate (vs. acute) calyx lobes, and from A. ericoides in its larger leaves. Sleumer (1959) separated the plants lacking gland-headed hairs as Leucothoé intermedia, but these plants are identical in all other characters to the glandular- hairy plants, and both pubescence forms can be found in the same locality (population?) (see Widgren 329 (Br, s), Widgren 668 (c, NY, $s), Lindberg 418 (ny, s), Leoncini & Roppa 341 (1); Regnell I-186 (F, M, NY, S, US), Regnell III- 838 (s), Regnell III-839 (s), Emmerich 2140 (L)). 22. Agarista ericoides Taubert, Bot. Jahrb. 17: 512. 1893. Ficure 7, d. Leucothoé ericoides (Taubert) Glaz. Bull. Soc. Bot. France 57(Mém. 3): 430. 1910. - Brazil, Minas Gerais, Serra dos Cristais, near Diamantina, 1892, Glaziou eT, ee jaca fragment of holotype, F!; photos of holotype, F!, Gu!; Res), Gi, Kl Pl), ae nee Sleumer, Bot. Jahrb. 78: 450. 1959. Type: Brazil, Minas Gerais, Serra de Ibitipoca (43°53’, 21°40’), 11 Aug. 1896, Schwacke 12360 (holotype, p!; photo of holotype, s!). Leucothoé acicularis Sleumer, ibid. 451. Tyre: Brazil, Minas Gerais, without definite locality, 1816-1821, St.-Hilaire, Cat. B', n. 2986 (holotype, P!; isotype, P!). Erect shrub to ca. 0.5(?) m tall. Twigs with or without scattered gland-headed hairs, otherwise glabrous to densely pubescent, with nonchambered pith. Buds to ca. 0.5 mm long. Leaves alternate; petiole 0.5—1.5 mm long, with or without gland-headed hairs adaxially; blade revolute in bud, narrowly ovate to nearly linear, 0.4-1.2 by 0.1-0.25 cm, strongly abaxially curved, coriaceous, the apex minutely acuminate, the base slightly cordate to truncate, the margin entire (to appearing serrulate/ciliate due to presence of many gland-headed hairs), strong- ly revolute, the adaxial surface with or without gland-headed hairs on extreme proximal portion of midvein, otherwise glabrous to very sparsely pubescent on proximal portion of midvein, the abaxial surface with or without gland- headed hairs along midvein, otherwise glabrous to very sparsely pubescent on extreme proximal portion of midvein, lacking or with few very inconspicuous glandular dots along midvein. Inflorescences axillary racemes to 1.5-3 cm long, the axis with or without scattered gland-headed hairs, otherwise densely pu- bescent. Pedicels 3.5-11 mm long, with or without gland-headed hairs, oth- erwise densely pubescent; bracteoles 2, alternate to opposite, from within lower ¥; to near middle of pedicel, narrowly triangular to ovate, to ca. 3 mm long; bracts ca. 3.5 mm long. Calyx lobes narrowly triangular, with long-acuminate apices, 3-6 by 0.4-0.7 mm, the abaxial surface with or without conspicuous gland-headed hairs, otherwise sparsely to densely pubescent, corolla cylindrical, 8-9.5 by 3.5-4 mm, pink to red, abaxially glabrous; filaments ca. 5.7-6 mm long, anthers ca. 1.5 mm long; ovary densely pubescent. Capsules short-ovoid, 3.5-4.5 by 4-5 mm, placentae + central; seeds 0.7-1.2 mm long. DISTRIBUTION. Southeastern Brazil, Minas Gerais (Map 7). REPRESENTATIVE SPECIMENS. Known only from type collections. This distinctive but poorly collected species is immediately recognizable due to its small, extremely revolute leaves. It is probably most closely related to 328 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Ficure 8. a, Agarista niederleinii var. niederleinii (Reitz & Klein 5889). b, A. nie- derleinii var. acutifolia (Hatschbach & Guimardes 25598). c, A. boliviensis (Steinbach 8568): note crisped/undulate leaf margins. . A. eucalyptoides (Williams & Assis 7503): note elongate, flexuous petioles. Scale = 2 ¢ 1984] JUDD, AGARISTA 329 three species from which it is geographically separated: Agarista hispidula, A. organensis, and A. chlorantha. Like many other species of Agarista, this taxon is quite variable in glandular indumentum. Some plants are densely covered with multicellular gland-headed hairs on stems and leaves, while others have only a few of these hairs or none at all. This variation thus cannot be used to delimit species, and Leucothoé lycopodioides and L. acicularis of Sleumer (1959) are reduced to synonymy under A. ericoides. 23. Agarista niederleinii (Sleumer) Judd, comb. nov. Shrub to small tree to 5 m tall with longitudinally furrowed bark. Twigs nearly glabrous to moderately pubescent, with nonchambered to clearly cham- bered pith. Buds to ca. | mm long. Leaves alternate; petiole 1.5-8 mm long; blade revolute in bud, ovate to elliptic or oblong, 0.6-5.7 by 0.3-1.7(-1.9) cm, flat to moderately abaxially curved, coriaceous, the apex acuminate or acute- to retuse-mucronate, the base cuneate to rounded (rarely very slightly cordate), the margin entire, plane to revolute, the adaxial surface glabrous, but sparsely pubescent on midvein, the abaxial surface glabrous (very sparsely pubescent along proximal portion of midvein), usually with few very inconspicuous glan- dular dots along midvein. Inflorescences axillary racemes to 1-3.5(-4.5) cm long, axis moderately to densely pubescent, bracteoles 2, alternate to opposite, from nearly basal to near midpoint of pedicel, triangular to linear, to ca. 1.5 mm long; bracts to ca. 1.7 mm long. Calyx lobes triangular with acuminate apices, 0.8-2.7 by 0.6-1.8 mm, abaxial surface glabrous to moderately pubes- cent; corolla cylindrical, 5-8 mm long, 2-4 mm wide, white, abaxially glabrous; filaments 4-5 mm long, anthers 0.9-1.5 mm long; ovary glabrous to sparsely pubescent (with few hairs at apex and around base). Capsules short-ovoid to subglobose, 4.5-7 by 5.5-10 mm, very thick walled, placentae subapical; seeds 1.5-3.2 mm long. DISTRIBUTION. Southeastern Brazil, from Parana south to Rio Grande do Sul in the Serra do Mar. Various habitats; 350-2000 m alt. Key TO THE VARIETIES OF AGARISTA NIEDERLEINII —_ Leaves with blade 0.6-2.8 by 0.3-0.9 cm, apex usually obtuse- to rounded- or retuse- mucronate; petiole 1.5-4.5 mm long. ..................05. a. var. niederleinii. 1. Leaves with blade 2-5.7 by 0.5—1.7(-1.9) cm, apex usually acute-mucronate to acu- iINinate: pevole 3=8 Mm log, 4.44640 sowie vane ee ee Peeks 23b. var. acutifolia. 23a. Agarista niederleinii (Sleumer) Judd var. niederleinii FIGURE 8, a. Leucothoé niederleinii Sleuamer, Notizbl. Bot. Gart. Berlin 12: 480. 1935. Type: Brazil, Santa Catarina, Campos de los Rios Chopim y Chapecé, Palmas Altas, Jan. 1887, Niederlein 2006 (holotype, B (destroyed)). The following neotype is here designated: Santa Catarina, Monte Cristo, Garuva, S. Francisco do Sul, Reitz & Klein 5889 (s!; isoneotypes, B!, L!, NY!, Uc!, Us!). 330 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 paeereteers . Map 8. Distribution of Agarista niederleinii var. niederleinii (dots), A. niederleinii var. acutifolia (circles), and A. u/eana (square). Leaves with petiole 1.5—4.5 mm long; blade 0.6-2.8 by 0.3-0.9 cm, flat to moderately abaxially curved, apex usually obtuse- to rounded- or retuse-mu- cronate, margin plane to revolute. Inflorescences to 1.5—3.5(—4.5) cm long; bracteoles narrowly triangular to linear. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, from Paranda south to Rio Grande do Sul in Serra do Mar (Map 8). Cloud forests, thickets, and open, rocky crests and slopes; 850-2000 m alt. Flowering November and December (to February). REPRESENTATIVE SPECIMENS. Brazil. PARANA: Guaratuba, Serra de Aracatuba, Hatschbach 6493 (L, UC). RIO GRANDE DO Suc: Taimbesinho, Araujo 1279 (1); S. Francisco de Paula, Cambara, Rambo 36724 (Ny); Serra da Rocinha, Bom Jesus, Rambo 53824 (B). SANTA CATARINA: Sao Joaquim, Serra do Oratorio, Lourteig 2149 (c, Pp, Ss, Us); Bom Retiro, Campo dos Padres, Reitz 2338 (B, L, NY, Uc, Us); Curral Falso, Bom Jardim, Sao Joaquim, Reitz & Klein 7794 (L); Morro do Campo Alegre, S. Francisco do Sul, Reitz & Klein 10322 (L, us); Campo Alegre, Morro Iquererim, Smith & Klein 8522 (1, us); Sao Joaquim, Serra do Oratério, 10 km E of Bom Jardim da Serra, Smith & Reitz 10159 (L, US). 23b. Agarista niederleinii (Sleumer) Judd var. acutifolia Judd, var. nov. IGURE 8, b. Varietas haec ab Agarista niederleinii var. niederleinii differt in foliis gran- dioribus 2-5.7 cm longis, 0.5-1.7(-1.9) cm latis, apicibus plerumque acumi- natis vel acutatis, mucronibus brevibus, et petiolis longioribus 3-8 mm longis. 1984] JUDD, AGARISTA 33% Leaves with petiole 3-8 mm long; blade 2-5.7 by 0.5-1.7(-1.9) cm, + flat, apex usually acuminate to acute- (obtuse-)mucronate, margin plane to very slightly revolute at base. Inflorescences to 1-2.8 cm long; bracteoles triangular. Type. Brazil, Parana, Campina dos Tavares, Bocaitiva do Sul, 21 Nov. 1970, G. Hatschbach & O. Guimardes 25598 (holotype, us!; isotypes, c!, M!, Mo!, ny!, s!, sp!, uc!). DISTRIBUTION AND ECOLOGY. Southeastern Brazil, from Paranda south to Rio Grande do Sul in Serra do Mar (Map 8). Thickets and thicket margins adjacent to grassy campo, stream margins; 350-1000(?) m alt. Flowering November and December. R Brazil. PARANA: Rio Pequeno, Sao José dos Pinhais, Hatsch- i 22818 (L, uc, Us); Campina dos Tavares, Bocaitiva do Sul, Hatschbach 23443 (L, y, us); Guaricana, Sao José dos Pinhais, Hatschbach 34903 (Ny). RIO GRANDE DO SUL: Taimbesinho, S. Francisco de Paula, Rambo 5412] (s). SANTA CATARINA: Morro Spitz- kopf, Klein 2318 (L, us), Reitz 2260 (ny, us); Alto Matador, Rio do Sul, Reitz & Klein 8303 (L Agarista niederleinii is most closely related to A. uleana, a geographically isolated species of the Pico da Tijuca region in Guanabara. Both species have distinctive, large, thick-walled capsules with subapical placentae, but they can be distinguished by leaf shape and size, and the extent to which the margins are revolute. The species is also easily confused with 4. minensis and A. pul- chella, both of which have smaller, thinner-walled capsules with usually more or less central placentae. In addition, A. pulchella has cordate-based leaves and usually longer racemes. It is of interest that Sleumer’s (1959) concept of A. niederleinii was quite broad, including all of the plants here referred to A. minensis and even a few here considered to belong to 4. pulchella. Although these three species are quite similar in leaf shape and size, they cannot be maintained as a single species because of the great variation in reproductive structures (especially inflorescence and fruit). Populations of Agarista niederleinii are separable into two morphologically distinctive and more or less elevationally isolated varieties. Variety niederleinii can usually be separated from var. acutifolia by its smaller, usually more or less obtuse-mucronate (vs. acute to acuminate) leaves. Variety niederleinii, from the specimens now available, seems to occur at higher and/or more exposed sites than var. acutifolia, and it is thus likely that the two taxa are ecologically isolated. However, the two may occasionally grow in close prox- imity, as they do near Taimbesinho, S4o Francisco de Paula, and a few more or less intermediate specimens (e.g., Rambo 54523, B) are known. These in- termediate plants appear to be highly fertile (most pollen grains stain darkly with cotton blue in lactophenol). Rambo 49305 (s; collected in Taimbe, S40 Francisco de Paula, Rio Grande do Sul) contains several somewhat aberrant twigs with small, more or less flat, cordate-based leaves. Sleumer (note on sheet) suggested that these plants pos- sibly represent hybrids between Agarista niederleinii (var. niederleinii) and A. pulchella var. cordifolia (a taxon with small, cordate-based leaves). However, A. pulchella var. cordifolia is limited to Minas Gerais and Sao Paulo and thus O52 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 65 does not occur anywhere near the state of Rio Grande do Sul. These plants are completely fertile (pollen grains stain darkly with cotton blue in lactophenol) and more likely represent merely a cordate-leaved extreme of A. niederleinii var. niederleinii. Field studies are needed in order to clarify this situation. 24. Agarista uleana (Sleumer) Judd, comb. nov. Leucothoé uleana Sleumer, Notizbl. Bot. he Berlin 12: 481. 1935. Type: Brazil, Guanabara, Rio de Janeiro, Pico da Tijuca, 1000 m alt., Nov. 1897, Ule 4576 ice la B (destroyed); isotype at HBG! ee designated as lectotype; isolectotype, Ee ambigua Meissner var. parvifolia Meissner in Martius, Fl. Brasil. 7: 156. 3. Type: Brazil, Guanabara, Rio de Janeiro, Riede/ s.n. (holotype, Herb. Sonder; fragment of holotype, Ny!). Shrub to small tree to 3 m tall. Twigs very sparsely to moderately pubescent, with nonchambered to clearly chambered pith. Buds to ca. 1.3 mm long. Leaves alternate; petiole 4-10 mm long; blade revolute in bud, ovate to elliptic (rarely oblong), 2-6 by 0.6-2.1 cm, + flat, coriaceous, the apex acuminate, to acute with small mucro, the base cuneate to rounded, the margin entire, very slightly revolute (+ plane), the adaxial surface glabrous, but sparsely pubescent on proximal portion of midvein, the abaxial surface glabrous (but sparsely pu- bescent along midvein), usually with few inconspicuous glandular dots along midvein. Inflorescences axillary racemes to 1-2.5(—4) cm long, axis moderately to densely pubescent. Pedicels 3-8 mm long, sparsely to densely pubescent; bracteoles 2, alternate to + opposite, nearly basal, triangular to narrowly so, to ca. 1 mm long; bracts to ca. 1.5 mm long. Calyx lobes triangular with acuminate apices, 1-2 by 0.6-1.2 mm, abaxial surface glabrous to moderately Sacao corolla cylindrical, 6.5-10 by 2—4.5 mm, white, abaxially glabrous; filaments 4.5-5.2 mm long, anthers |-1.6 mm long; ovary glabrous to sparsely pubescent, especially near apex and base. Capsules short-ovoid to subglobose, (3-)4-6 by (5-)5.5-8 mm, very thick walled, placentae subapical; seeds 1.5- 2.8 mm long. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, Guanabara, chiefly limited to Pico da Tijuca region (Map 8). Low forest and scrub, sunny open areas; ca. 700-1020 m alt. Flowering September through November REPRESENTATIVE SPECIMENS. Brazil. GUANABARA: Pico da Tijuca, A Araujo & Almeida 519 (F), Duarte 8462 (L, M, NY, 8, US); Morro Queimado, Brade 11108 (Gu), Glaziou 6620 (c, F, NY, P, 8); Pedra do Conde, Tijuca, Lems s.n., 22 March 1964 (F, L, M, MO, NY, 8). Estapo bo Rio: see Sleumer (1959). Agarista uleana is most closely related to A. niederleinii and is especially easily confused with 4. niederleinii var. acutifolia. Both species have distinc- tive, large, thick-walled capsules with subapical placentae. Agarista uleana can be separated from A. niederleinii var. niederleinii by its much larger leaves with acute to acuminate (vs. obtuse- to rounded-mucronate) apices, and from var. acutifolia by its slightly different range of leaf shapes and by the extent to which its leaf margins are revolute. The two taxa are geographically separated. Nonfruiting specimens may be confused with A. o/eifolia, A. populifolia, or A. 1984] JUDD, AGARISTA 330 mexicana, but none of these species occurs in the Pico da Tijuca region; see key for distinguishing characters. 25. Agarista boliviensis (Sleumer) Judd, comb. nov. FIGURE 8, Cc. Leucothoé boliviensis Sleumer, Notizbl. Bot. Gart. Berlin 12: 131. 1934. Type: Bolivia, Dept. Santa Cruz, valley of Comarapa, 2000 m alt., 26 Oct. 1928, Steinbach 8568 (holotype, B (destroyed); fragment of holotype, us!; isotypes, BM, el, F!, G, GH!, K!, LIL, Mo!, Ny!, s!, uc!; photos of isotype, F!, G, GH!). Shrub or small tree to ca. 5 m tall. Twigs glabrous, with obscurely chambered pith. Buds to ca. 1 mm long. Leaves alternate; petiole (5-)8-17 mm long, frequently slender and flexuous; blade revolute in bud, ovate, 2-5.5 by 0.9- 2.2 cm, + flat, coriaceous, the apex acuminate, the base rounded and often slightly asymmetric, the margin entire and minutely undulate, + plane, the adaxial surface glabrous, but usually very sparsely pubescent on midvein, the abaxial surface glabrous, but usually very sparsely pubescent along midvein, with or without inconspicuous to conspicuous glandular dots along midvein. Inflorescences axillary racemes to 1-2.5 cm long, axis moderately pubescent with + whitish hairs. Pedicels 4-8 mm long, sparsely to moderately pubescent; bracteoles 2, opposite to alternate, from basal to within lower ' of pedicel, narrowly triangular to linear, to ca. 1.1 mm long; bracts to ca. 1.4 mm long. Calyx lobes triangular with acuminate apices, 0.9-1.7 by 0.5-1.5 mm, abaxial surface glabrous to moderately pubescent; corolla cylindrical, 6-9.5 by 3-5 mm, white, abaxially glabrous; filaments 3.5—4.5 mm long, anthers 1-1.1 mm long, ovary glabrous to sparsely pubescent near apex. Capsules subglobose to ovoid, 3.5-4.5 by 5-6.5 mm, placentae subapical; seeds 2-2.5 mm long DISTRIBUTION AND ECOLOGY. Bolivia (Map 9). Mountainous areas ca. 1200- 2500 m alt. Flowering September and October. REPRESENTATIVE SPECIMENS. Bolivia: Charcas, San Pedro, Pasopaya, Anonymous 3400 peladas, Alto de las Cafias, Troll 359 (g, mM); Camino de Emborozt,, La Mamora, Ttirpe et al. 4777 (BAA). Agarista boliviensis is most closely related to A. eucalyptoides, from which it is easily distinguished by its more consistently ovate leaves and its inflores- cence axis with whitish (vs. ferrugineous) hairs. Both taxa are distinctive due to their moderate-sized leaves with often elongate and flexuous petioles. Agar- ista boliviensis has consistently crisped/undulate leaf margins, whereas in A. eucalyptoides this character is variable. The two species are completely allo- patric since A. eucalyptoides is limited to southeastern Brazil and Uruguay. 26. Agarista eucalyptoides (Cham. & Schldl.) G. Don, Gen, Syst. 3: 837. 1834. FiGure 8, d. Andromeda lanceolata Vell. Conc. Fl. Flum. 175. 1825, later homonym of A. lanceolata Wallich, Asiatic Res. 12: 391. 1820 = Lyonia ovalifolia (Wallich) Drude. TyPe: “Cabinet d’Histoire naturelle de Rio de Janeiro” (not seen 334 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Map 9. Distribution of Agarista boliviensis (circles), A. eucalyptoides (dots), A. gla- berrima (triangles), A. angustissima (open squares), and 4. duartei (closed squares). Andromeda eucalyptoides Cham. & Schlidl. Linnaea 1: 518. 1826. Leucothoé eucalyp- toides (Cham. & Schldl.) DC. Prodr. 7: 605. 1839. res multiflora (Pohl) DC. var. Benne (Cham. & Schldl.) Meissner in Martius, Fl. Brasil. 7: 155. 1863. Type: Brazil, Rio Grande do Sul, Sellow s.n. floc B (destroyed); fragment of holotype, Ny; isotypes, BR, E!, G!, kK! oe multiflora Pohl, PI. Brasil. 2: 33. t. 122. 1828/29. Agarista multiflora ohl) G. Don, Gen. Syst. 3: 837. 1834. Leucothoé multiflora (Pohl) DC. Prodr. 7: ie 1839. Leucothoe multiflora (Pohl) DC. var. pohlii Meissner in Martius, Fl. Brasil. 7: 155. 1863, nomen superfl. Tyre: Brazil, Minas Gerais, Rancho Novo, seg ae Mantiqueira, Sept./Oct. 1819, Pohl s.n. (holotype, w (destroyed?); isotypes, Andiomet longepetiolata Fenzl ex Ettingsh. Blatt-Skel. Dikot. 94. fig. 56. 1861. Type: fhevebinoe multiflora (Pohl) DC. var. petiolaris Meissner in Martius, Fl. Brasil. 7: 155. 863 zil, Minas Gerais, Serra de Cural d’El Rey, ca. 1841-1842, Gardner 498 86 (ho en BM; isotypes, E!, F!, G!, GH!, K!, _ p!, us!, w). Leucothoé ie (Pohl) DC. var. eines Meissner in Martius, ibid. Tyre: Brazil, Minas Gerais, Tejuco (= Diamantina), Martius 1337 (holotype, M!; isotype, M!). Shrub or small tree to ca. 4(—8) m tall, with often contorted trunks and thick, corky, furrowed bark. Twigs glabrous to sparsely pubescent, with + noncham- bered pith. Buds to ca. 1.5 mm long. Leaves alternate to subopposite or + whorled, often variable even within single twig; petiole 6-40 mm long, fre- quently slender and flexuous; blade revolute in bud, ovate to oblong, 2.5-8 (-9.3) by 0.8-2.6 cm, + flat, coriaceous, the apex acute to acuminate or shortly 1984] JUDD, AGARISTA 335 so, to nearly rounded-mucronate, the base rounded to truncate and often slightly asymmetric, the margin entire (undulate), plane to very slightly revolute, es- pecially near base, the adaxial surface glabrous (sparsely pubescent on midvein), the abaxial surface glabrous (very sparsely pubescent on midvein near base), with usually at least few inconspicuous (to conspicuous) glandular dots along midvein. Inflorescences axillary racemes to 1-6 cm long, axis moderately to densely ferrugineous/crisped-pubescent. Pedicels 2-6 mm long, sparsely to densely ferrugineous-pubescent; bracteoles 2, opposite to alternate, from nearly basal to near midpoint of pedicel, narrowly triangular, to ca. 1.1 mm long; bracts to ca. 1.7 mm long. Calyx lobes triangular with acuminate (to acute) apices, 1-1.8 by 0.5-1.2 mm, abaxial surface essentially glabrous, corolla cy- lindrical, 6-10.5 by 2.5-5 mm, white (to reddish), abaxially glabrous; filaments 3-4.5 mm long, anthers 0.8-1.1 mm long; ovary glabrous to sparsely pubescent. Capsules ovoid to short-ovoid, 3-5 by 4-5 mm, placentae subapical; seeds 1.8-2.5 mm long. DISTRIBUTION AND ECOLOGY. Southeastern Brazil (Minas Gerais and adjacent regions of Estado do Rio and Sao Paulo; Santa Catarina, Rio Grande do Sul) and Uruguay (Map 9). Cerrado vegetation, open, rocky thickets and thicket margins, or rocky hillsides; (250-)800-2300 m alt. Flowering chiefly late August through November (December). REPRESENTATIVE SPECIMENS. Brazil. Estapo po Rio: Nova Friburgo to Pedra do Cénego, Glaziou 12927 (c, Pp). Minas Gerais: Itatiaia, Brade 14086 (F, L); Serra do Caparao, Brade 16987 (F, L); So Thomé das Letras, Baipendi, Brade 20429 (F, m); Serra do Cipd, m 131, Duarte 2693 (r, L); Serra da Moeda, Duarte 8897 (L, M); Serra do Itatiaia, Dusén 2011 (G, GH, s); Serra da Mutuca, Belo Horizonte, Markgraf 3523 (F); 35 km from Lambari, Pereira 7153 (s), Uberava, Regnell III-837 (s, us); Pico da Bandeira, near Caparao, Shepherd et al. 5795 (1); Caldas, Widgren 327 (s);, Serra da Calveira, Betim, Williams & Assis 7503 (F, GH, MO, NY, S, SP, US). RIO GRANDE DO SUL: Porto Alegre, Morro da Policia, Ma/lme 608 (s); Sao Leopoldo, Rambo 132 (s, sp); Montenegro, Zim- merberg, Rambo 8309 (B); S. Francisco de Paula, Vila One, alla 31063 (F, MO, S); Pérto Alegre, Morro da Gloria, Rambo 40072 (B, Mo); P rro, Vacaria, Rambo 51626 (Ss, US). SANTA CATARINA: 16 km E of Lajes on rd. to Painel, Smith & Reitz 10103 (L, us). SAo PauLo: Campos do Jordao, Leite 3614 (A, GH). eee. Rivera, Cufiapird, Berro 4973 (a); Rivera, Galgo, Herter 1859 (8, F, G, L, M, MO, Agarista eucalyptoides is a distinctive species easily recognizable by the mod- erately to densely ferrugineous/crisped pubescence of its inflorescence axis and pedicels. It is most closely related to A. boliviensis, a geographically separated species with more consistently ovate leaves and an inflorescence axis with whitish hairs. Both of these species (along with A. glaberrima and—to a lesser extent—A. angustissima) frequently have slender, flexuous, elongate petioles. Sleumer (1959) considered 4. glaberrima to be only varietally distinct from A. eucalyptoides, but the two taxa differ in leaf shape, degree of adaxial folding of the lamina, inflorescence indumentum (lacking in A. glaberrima), and—to a lesser extent—capsule size. The morphological gap separating these taxa 1s thus comparable to that between most Agarista species. Agarista glaberrima is considered here to be a distinct species that is probably more closely related to A. angustissima than it is to A. eucalyptoides. 336 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 bint m=. ey a, Agarista glaberrima (Hatschbach, Smith, & Ayensu 28814). b, A. an- m. FIGURE 9. a, gustissima (Hatschbach 27988, Uc): note terminal racemes. Scale = 2 c 27. Agarista glaberrima (Sleumer) Judd, comb. et stat. nov. FIGURE 9, a. Leucothoé eucalyptoides (Cham. & Schldl.) DC. var. glaberrima Sleumer, Bot. Jahrb. 78: 458. 1959. Type: Brazil, Minas Gerais, Santa Lucia, Serra do Cipé, km 135, 4 Feb. 1938, Barreto 8921 (holotype, F!; photo of holotype, F'). Shrub or small tree to 4 m tall. Twigs glabrous, with nonchambered to clearly chambered pith. Buds to ca. | mm long. Leaves alternate; petiole 10-32 mm long, frequently slender and flexuous; blade revolute to conduplicate(?) in bud, ovate to narrowly ovate, 2.3-8.5 by 0.6-2.5(-3) cm, flat to strongly adaxially folded, coriaceous, the apex acuminate, the base cuneate to rounded and often slightly asymmetric, the margin entire (slightly undulate), plane, the adaxial surface glabrous (very sparsely pubescent on proximal portion of midvein), the abaxial surface glabrous, with few to many inconspicuous to conspicuous glan- dular dots along midvein. Inflorescences axillary racemes, or terminal racemes or panicles, to 1-9.5 cm long, axis glabrous. Pedicels 2.5-8 mm long, glabrous; bracteoles 2, opposite to alternate, from nearly basal to near midpoint of pedicel, narrowly triangular, to ca. 0.9 mm long; bracts to ca. 1.4 mm long. Calyx lobes triangular with acuminate (to nearly acute) apices, 0.8-1.7 by 0.5- 1.3 mm, abaxial surface glabrous; corolla + cylindrical, 6-10 by 2.7-4.5 mm, white or greenish white, abaxially glabrous; filaments 3.5—4.5 mm long, anthers 1.1-1.3 mm long; ovary glabrous or nearly so. Capsules subglobose to short- ovoid, 4-8 by 6-8 mm, placentae subapical to nearly central; seeds 2—2.5 mm long. 1984] JUDD, AGARISTA ao7 DISTRIBUTION AND ECOLOGY. Southeastern Brazil, Minas Gerais (Map 9). Sandy to rocky cerrado vegetation, margins of woods, rocky and/or disturbed thickets, rocky hillsides; 900-1500 m alt. Flowering chiefly November through February. REPRESENTATIVE SPECIMENS. Brazil. MinAs GERAIs: Serra do Espinhaco, Gouvéa, km 258 on M. G. 259, Anderson et al. 35627 (F, L, Ny); Serra dos Cristais, Diamantina, Barreto 9562 (F); Boa Vista-Extraccéo, Diamantina, Barreto 9647 (F); Serra do Cabral, Joaquim Felicio, Davis et al. 2374 (cE); Diamantina, Duarte 10526 (L); Serra do Cipé, araalear Hatschbach et al. 28814 (c, L, MO, NY, 8, UC); Serra do Espinhaco, Lapinha, 19 km N Cérro on road to Diamantina, /rwin et al. 20802 (Ny), 12 km NE of Diamantina on rd. to Mendanha, Irwin et al. 22758 (ENCB, F, GH, K, NY); Serra Grao Mogul, Maguire et al. 49269 (ny); Serra Negra, St.-Hilaire, Cat. D, n. 94 (L, P). Agarista glaberrima is most closely related to the sympatric A. angustissima, another Ce ae endemic (Map 9) from which it can easily be distin- guished by vate and wider leaves with usually longer petioles and less strongly aed folded blades. Although Sleumer (1959) considered this species to be a variety of A. eucalyptoides, the two taxa are quite distinct and can be separated by inflorescence indumentum (lacking vs. moderate to dense and ferrugineous), leaf shape (ovate vs. ovate to oblong), and degree of adaxial folding of the lamina. 28. Agarista angustissima Taubert, Bot. Jahrb. 17: 513. 1893. Ficure 9, b. Leucothoé angustissima (Taubert) Sleumer, Bot. Jahrb. 78: 451. 1959. Type: Brazil, Minas Gerais, Pinheiro, near Biribiri, 26 March 1892, Glaziou 19582 (holotype, B (destroyed); fragment and photo of holotype, F!; photo of holotype, Gu!; isotypes, BR, F!, G!, K!, Mo!, NY!, P!). Shrub to ca. 2 m tall. Twigs with or without gland-headed hairs, otherwise glabrous, with nonchambered pith. Buds to ca. 1 mm long. Leaves alternate; petiole 5.5-13 mm long, often slender and flexuous; blade conduplicate in bud, linear and curved, to narrowly ovate in juvenile leaves, (1.7—)3-7 by 0.1-0.5 cm but often appearing narrower, strongly adaxially folded and thus obscuring adaxial surface, coriaceous, the apex narrowly acute to short-acuminate, the base narrowly cuneate, the margin entire, to serrulate in juvenile leaves due to presence of gland-headed hairs, plane, the adaxial surface glabrous, the abaxial surface with or without gland-headed hairs, otherwise glabrous, lacking or with few inconspicuous glandular dots along midvein. Inflorescences axillary ra- cemes, or terminal racemes or panicles, to 3-6 cm long, axis glabrous. Pedicels 4.5-14 mm long, glabrous; bracteoles 2, alternate to opposite, from nearly basal to within lower '4 of pedicel, narrowly triangular to linear, to ca. | mm long; bracts to ca. 3 mm long, grading into leaves. Calyx lobes triangular with acu- minate apices, 1-1.5 by 0.5-1.1 mm, abaxial surface glabrous; corolla cylin- drical, 6.5-10.5 by 2.5-3.5 mm, white, abaxially glabrous; filaments 4-5 mm long, anthers 1-1.7 mm long; ovary glabrous. Capsules short-ovoid to sub- globose, 3.5-5.5 by 5-7 mm, placentae subapical to nearly central; seeds 1.5- 2.9 mm long. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, Minas Gerais (MAP 9). Ce- rrado vegetation, moist rocky areas; ca. 1200-1400 m alt. Flowering November. 338 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 REPRESENTATIVE SPECIMENS. Brazil. MINAS Gerats: Diamantina, Barreto 10132 (F); Biri- biri, Diamantina, Hatschbach & Pelanda 27988 (c, L, NY, 8, UC); Serra do Cip6, Fazenda Sue Jaboticatuba, Hatschbach & Ahumada 31564 iL) Chapada, St.-Hilaire, Cat. , A. 2057 (L, P). Agarista angustissima 1s a very distinctive species recognizable by its very narrow, more or less linear/curved leaves that are strongly adaxially folded and ca. 3-7 cm long, its glabrous stems, and its often terminal inflorescences. Most closely related to 4. glaberrima and A. duartei, it can be distinguished from the former by its narrower, linear leaves, and from the latter by its glabrous stems and its more elongated, terminal inflorescences. All three species are limited to the Serra do Espinhaco region of Minas Gerais. 29. Agarista duartei (Sleumer) Judd, comb. nov. Leucothoé duartei Sleumer, Bot. Jahrb. 78: 451. 1959. Type: Brazil, Minas Gerais, Serra do Cipé, km 137 da Estrada de Concei¢ao, 1300 m alt., 6 Dec. 1949, Duarte 2103 (holotype, RB; isotypes, L!, LIL). Subshrub to ca. 0.5 m tall, with bark not well developed. Twigs + glabrous to densely pubescent, with nonchambered to irregularly chambered pith. Buds to ca. | mm long. Leaves alternate; petiole 2-6.5 mm long, not flexuous; blade conduplicate in bud, linear and + curved, 1.2—3.5(-4) by 0.2-0.4 cm but often appearing narrower, strongly adaxially folded and thus obscuring adaxial sur- face, coriaceous, the apex narrowly acute to short-acuminate, the base narrowly cuneate, the margin entire, to serrulate in juvenile leaves due to presence of gland-headed hairs, plane, the adaxial surface sparsely pubescent, the abaxial surface with or without gland-headed hairs, otherwise glabrous to very sparsely pubescent on proximal portion of midvein, lacking or with few inconspicuous glandular dots along midvein. Inflorescences axillary racemes to 0.3-1.5 cm long, axis very slightly to densely pubescent. Pedicels 2-7.5 mm long, glabrous to densely pubescent; bracteoles 2, subopposite to alternate, from nearly basal to near midpoint of pedicel, narrowly triangular, to ca. 1 mm long; bracts to ca. 1 mm long. Calyx lobes triangular with acuminate apices, 1-1.5 by 0.6-1 mm, abaxial surface essentially glabrous; corolla cylindrical, 7-8 by 2.5-3.5 mm, white, abaxially glabrous; filaments 4.5—5 mm long, anthers 1.1-1.2 mm long; ovary glabrous to moderately pubescent, placentae + subapical(?). Cap- sules not seen. DISTRIBUTION AND ECOLOGY. Southeastern Brazil, Minas Gerais (Map 9). Open sandy or rocky areas; ca. 1300 m alt. Flowering October to December REPRESENTATIVE SPECIMEN. Brazil. Minas Gerais: Serra do Cipé, Sta. Ana do Riacho, Hatschbach & Koczicki 35379 (1). This rare and little-collected species is most closely related to Agarista an- gustissima and A. glaberrima. It is easily distinguished from both of these species by its often pubescent stems and inflorescence axis, its shorter petioles, and its shorter and exclusively axillary racemes. Its leaves, like those of A. angustissima, are linear and very strongly adaxially folded. Agarista duartei 1984] JUDD, AGARISTA 339 has only been collected in the Serra do Cipé, a region where the genus shows a large amount of diversity. Fruiting material has not been seen. Agarista D. Don ex G. Don sect. Agauria (DC.) Judd, comb nov. Leucothoé D. Don sect. Agauria DC. Prodr. 7: 602. 1839. Agauria (DC.) J. D. Hooker in siag erp i Pl. 2: 586. 1876. Lecrorype species: Agarista salicifolia (Comm. ex Lam.) G Twigs with nonchambered, slightly to very heterogeneous pith. Leaves with the abaxial epidermis papillose, the adaxial epidermal cells short, usually not divided. Style apparently not swollen. Capsules with placentae basal; seeds 3- 4 mm long. DistRiBUTION. Central Africa, Madagascar, Réunion, and Mauritius (see Sleu- mer, 1938, fig. /). NUMBER OF SPECIES (TAXA): | (20). This section is not treated here; see Sleumer (1938) for a revision of the group and a key to the many infraspecific taxa comprising the widespread and variable Agarista salicifolia (species no. 30). SPECIES EXCLUDED FROM AGARISTA Agarista anastomosans G. Don, Gen. Syst. 3: 838. 1834, nomen superfl. = Gaultheria glomerata (Cav.) Sleumer (Sleumer, 1959). Agarista ciliata (Nees) J. D. Hooker ex Niedenzu, Bot. Jahrb. 11: 236. 1889 = Gaylussacia brasiliensis (Sprengel) Meissner (Sleumer, 1959). Agarista coccinea (Schrader) J. D. Hooker ex Niedenzu, ibid. = Gaylussacia brasiliensis (Sprengel) Meissner (Sleumer, 1959). Agarista eriophylla (Pers.) G. Don, Gen. Syst. 3: 838. 1834 = Gaultheria eriophylla (Pers.) Sleumer ex Burtt (Sleumer, 1959). Agarista ilicifolia (Pers.) G. Don, Gen. Syst. 3: 838. 1834 = Gaultheria phil- lyreifolia (Pers.) Sleumer (Sleumer, 1959). Agarista itatiaiae (Wawra) Wawra, Itin. Princ. Coburgi 1: 73. 1883 = Gaul- theria itatiaiae Wawra (Sleumer, 1959). ACKNOWLEDGMENTS I wish to thank the curators of the herbaria from which specimens have been borrowed for this study (A, B, BAA, BAF, BM, C, COL, DUKE, E, ENCB, F, FLAS, GH, GOET, HBG, K, L, M, MEXU, MO, NA, NCU, NY, P, S, SP, UC, US, UWFP, VEN, WIS). I am particularly grateful to Kent D. bea registrar of the Oniversity of Florida Herbarium, for his help in obtaining and processing specimen loans, and Peter F. Stevens and James L. Luteyn for their many helpful suggestions concerning the manuscript. I also want to thank Stephen A. Spongberg and Elizabeth B. Schmidt for their helpfulness, corrections, and suggestions in con- verting this manuscript into print. 340 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 LITERATURE CITED BaRTRAM, W. 1791. Travels through North & South Carolina, Georgia, East & West Florida. 522 pp. James & Johnson, Philadelphia. BENTHAM, G., & J. D. Hooker. 1876. Leucothoé D. Don, Agauria DC., Agarista D. Don. Gen. Pl. 2(2): 584-586. CANDOLLE, A. P. bE. 1839. Waccinieae, Ericaceae. Prodr. 7: 522-733. Cox, H. T. 1948. Studies in the comparative anatomy of the Ericales II. Ericaceae - subfamily Arbutoideae. Am. Midl. Nat. 40: 493-516. Crisci, J. V., & T. F. Sruessy. 1980. Determining primitive character states for phy- logenetic reconstruction. Syst. Bot. 5: 112-125. Don, G. 1834. A general system of dichlamydeous plants. Vol. 3. 867 pp. J. G. & F. Rivington et a/., London Dresscer, R. L. 1954. Some floristic relationships between Mexico and the United States. Rhodora 56: 81-96. Drupe, O. 1897. Ericaceae. Jn; A. ENGLER & K. PRANTL, Nat. Pflanzenfam. IV. 1: -65. EIreEN, G. 1972. The cerrado vegetation of Brazil. Bot. Rev. 38: 201- E.uioTr, S. 1816-1821. Sketch of the botany of ee Carolina and Sk Vol. 1. 606 pp. J. R. Schenck, Charleston, South Car Emricu, K., & B. RAMBo. Florae Riograndensis. Lilloa 17: 106-108. GOMEz- PoMPa: A. 1973. Ecology of the vegetation of Veracruz. Pp. 73-148 in A. GRAHAM, ed., ees and vegetational history in northern Latin America. El- sevier Publ. Co., York. — R. Ee The peat of the flowering plants. ed. 3. 518 pp. Longman Group ondon. Sa 7 1964. Origin and evolution of the biota of a aia North America: evidence from the fossil plant record. Evolution 18: 571-585. 19 History of the arborescent temperate a in the northern Latin American biota. Pp. 301-314 in A. GRAHAM, ed., Vegetation and vegetational his- tory in northern Latin America. Elsevier Publ. Co., New York. Gray, A. 1878. Andromeda L. Synoptical flora of North America. ed. 2. Vol. 2, part 1, pp. 33-35. Ivison, Blakeman, Taylor and Co., New Yor Ha.té, F., R. A. A. OLDEMAN, & P. B. TOMLINSON. 1978. Tropical trees and forests: an architectural analysis. 441 pp. Springer-Verlag, Berlin. HoLMGREN, P. K., W. KEUKEN, & E. K. SCHOFIELD. 1981. Index herbariorum. ed. 7. 23 Hooker, W. J. 1847. Leucothoé pulchra. Curtis’s Bot. Mag. 73: t. 4314. & J. SmitH. 1851. Leucothoé neriifolia. Curtis’s Bot. Mag. 77: t. 4593. INGRAM, J. 1961. Studies in the cultivated Ericaceae 1. Leucothoé. tare 9: 57-66. Jupp, W.S. 1979. Generic relationships in the Andromedeae ( ). Jour. Arnold Arb. 60: 477-503. 1981. A monograph of Lyonia (Ericaceae). Ibid. 62: 63-209, 315-436. 82. A taxonomic revision of Pieris (Ericaceae). Ibid. 63: 103-144. KINosHITA-GouvEa, L. 1980. Estudos taxonomicos e fitogeograficos da familia Eri- caceae do Brasil. Unpubl. Ph.D. Thesis, Univ. SAo Paulo. LAMARCK, J. B. 1783. Encyclopédie méthodique. Vol. 1. 752 pp. Plomteaux, Paris. s, K. 1962. Adaptive radiation in the Ericaceae. I]. Shoot development in the Andromedeae. Ecology 43: 524-528. . 1964. Evolutionary studies in the Ericaceae. IJ. Leaf anatomy as a phylogenetic index in the Andromedeae. Bot. Gaz. 125: 178-186. LoESENER, T. 1889. Ueber einige neue Pflanzenarten aus Brasilien. Flora 72: 74-79. Macauireg, B., J. A. STEYERMARK, & J. L. LuTEyn. 1978. Ericaceae. Jn: B. MAGUIRE & 1984] JUDD, AGARISTA 34] COLLABORATORS aa botany of the Guayana Highland—part X. Mem. New York Bot. Gard. 29: -203. Marques, M. po C M., & R. M. Kren. 1975. Flora aha catarinense parte I: Ericaceas. 65 pp. Itajai, Herbario “Barbosa Rodrigue Martin, P. S., & B. E. HARRELL. 1957. The Pleistocene ee of temperate biotas in Mexico and the eastern United States. Ecology 38: oe MEISSNER, C. F. 1863. Leucothoé. In: C. F. P. von Martius, Fl. Brasil. 7: 154-166. Me vin, N.C. 1981. Additional comments on Leucothoé aac Baileya 21: 127- 30 130. MicHAux, A. 1803. Flora Boreali-Americana. Vol. 1. 330 pp. Levrault, Pari MIRANDA, F., & A. J. SHARP. 1950. Characteristics of the a a regions of eastern Mexico. Ecology 31: 313-333. NieEDENzu, F. 1889. Uber den anatomischen Bau der Laubblatter der Arbutoideae und Vaccinioideae in Beziehung zu ihrer systematischen Gruppierung und geographisch- en Verbreitung. Bot. Jahrb. 11: 134-263. NuttacL, T. 1818. Genera of North American plants. Vol. Philadelphia. Rzepowskl, J. 1965. ea oa y posibles origenes de la flora de México. Bol. Soc. Bot. México 29: -177. SLEUMER, H. 1938. Die tts Agauria (DC.) Hook. f. Bot. Jahrb. 69: 374-394. . 1959. Studien iiber die Gattung Leucothoé D. Don. Ibid. 78: 435-480. SMALL, J. K. 1914. Ericaceae. N. Am : 33-102. Smitu, A. C. 1950. Studies of South Ponce plants, XII. Contr. U. S. Natl. Herb. 29: 335, 336. Situ, L. B. 1962. Origins of the flora of southern Brazil. Contr. U.S. Natl. Herb. 35: 215-249. — 312 pp. D. Heartt, ee P.C., & L. O. WituiaMs. 1952. Leucothoé. Flora of Guatemala. Fieldiana t. 24: 108-110. curr P.F. 1970. Agauria and Agarista: an example of tropical transatlantic affinity. Notes Royal Bot. Gard. Edinburgh 30: 341-359. : rhe A classification of the Ericaceae: subfamilies and tribes. Bot. Jour. Linn. Soc. 64: ——.. ee : peenennay polarity of character states. Ann. Rev. Ecol. Syst. 11: 333- 1981. On ends and means, or how polarity criteria can be assessed. Syst. Bot. 8. TRON: R. 1972. Endemic areas and geographic speciation in tropical American ferns. Biotropica 4: 121-131. WAGNER, W. H. 1961. Problems in the classification of ferns. Pp. 841-844 in W. B. TurriLL, ed., Recent advances in botany. Vol. 1. Univ. Toronto Press, Toronto. 1962. A graphic method for expressing relationships based upon group cor- relations of indexes of divergence. Pp. 415-417 in L. Benson, Plant taxonomy: methods and principles. Ronald Press Co., New York 1 The construction of a classification. Pp. 67-90 in C. G. SIBLEy, ed., Systematic biology. National Academy of Science Publ 1692, Washington D.C. . Origin and philosophy of the g 2 cladistics Syst. ao 5: 173-193. Watrous, L. E., & QUENTIN D, WHEELER. 1981. The out-group comparison method of character analysis. Syst. Zool. 30: 1-11. WEBERLING, F. 1965. Typology of inflorescences. Jour. Linn. Soc. Bot. 59: 215-221. WHEELER, Q. D. 1981. The ins and outs of character analysis: a response to Crisci and Stuessy. Syst. Bot. 6: 297-306. Witey, E. O. 1981. Phylogenetics. 439 pp. John Wiley & Sons, New York. 342 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Woop, C. E. 1961. The genera of the Ericaceae in the southeastern United States. Jour. Arnold Arb. 42: 10-80. Wurback, J. 1970. Erroneous data in Glaziou collections of Melastomataceae. Taxon : 911-913. DEPARTMENT OF BOTANY FLORIDA GAINESVILLE, FLORIDA 3261 1 1984] AL-SHEHBAZ, CRUCIFERAE 343 THE TRIBES OF CRUCIFERAE (BRASSICACEAE) IN THE SOUTHEASTERN UNITED STATES! IHSAN A. AL-SHEHBAZ The family Cruciferae is represented in the southeastern United States by 121 species in 43 genera assigned to seven tribes. The present account includes a family description, with general comments on the group as a whole; selected family references; a key to the tribes; an artificial key to 46 genera (including three known as escapes from cultivation to the west of this area); and brief descriptions of the tribes, each with a list of the representative genera in the outheast. When treating a family generally recognized as difficult for generic and tribal delimitation, one faces the problem of how genera should be arranged. An alphabetic sequence would definitely be incompatible with the scope of our flora. A few students of the family avoid recognizing tribes because they believe that tribal boundaries are usually artificial. However, these authors arrange the genera according to their nearest sister relatives—a disposition that often co- incides so well with the tribal classification that ignoring or totally abandoning the tribes is unreasonable. Nearly half of the genera of Cruciferae occurring in our area belong to the tribes Thelypodieae, Brassiceae, and Lepidieae, which are widely recognized as natural groups. The majority of the remaining genera fall within the presumably well-defined centers of four other tribes. For these reasons I favor the use of tribes to provide a workable framework, even though ‘Prepared for fthe Soutt United States, a long-term project made possible . kane — the National Science Foundation and currently ee by BSR-8111520 (C. E. d, Jr. ipal investigator), under which this research was done, and BSR-8303100 (N. G. ie Seal investigator). This account, the 103rd in the series, follows the format established in the first paper (Jour. Arnold Arb. 39: 296- oe 1958) and continued to the present. The area covered by the Generic Flora includes North and South Carolina, Georgia, Florida, Tennessee, Alabama, Mississippi, Arkansas, and Louisiana. The descriptions are based primarily on the plants of this area, with information about ehleiceres members of a family or genus in brackets [ ]. The two references that I have not verified are marked with an asterisk. most grateful to Carroll Wood for tis support, guidance, and help with many aspects of this paper, and particularly for his critical review of the manuscript. Reed C. Rollins, Norton.G. Miller, George K. Rogers, and Elizabeth A. Shaw were helpful throughout this study, and I am indebted to Barbara Nimblett for her careful typing of the manuscript. I am grateful to Elizabeth B. Schmidt and Stephen A. Spongberg for their editorial help. The illustrations were made by Karen Stoutsenberger (primarily under earlier grants), with the exception of Figures 1, s (by Rachel A. Wheeler) and 3, a-c (by the late Dorothy H. Marsh). Carroll WwW seeds are largely from ieeecium specimens of the Arnold Arboretum and the Gray Herbarium. © President and Fellows of Harvard College, | Journal of the Arnold Arboretum 65: 343-373. ae 1984. 344 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 I fully admit that, when taken on a worldwide basis, some tribal boundaries become clearly artificial. CRUCIFERAE A. L. de Jussieu, Gen. Pl. 237. 1789. Nom. alt. BRASSICACEAE Burnett, Outlines Bot. 1123. 1835. (MUSTARD FAMILY) Annual, biennial, or perennial herbs [rarely vines, subshrubs, shrubs, or even small trees], with pungent, watery juice rich in glucosinolates (mustard-oil glucosides) and with idioblasts containing myrosinase; indumentum lacking or of simple, furcate, dendritic, or stellate unicellular, eglandular trichomes, very rarely with additional unicellular or multicellular glands. Leaves alternate [very rarely opposite], sometimes confined to a basal rosette, exstipulate, simple, very rarely pinnate or palmate, as in Cardamine; stomata typically anisocytic, rarely mixed with few of some other types. Inflorescences terminal (axillary in Coronopus), usually racemes, corymbs, or panicles, rarely solitary on long scapes (Leavenworthia), usually ebracteate (bracteate in Se/enia). Flowers hy- pogyn perfect]. Sepals 4, almost always free, in 2 decussate pairs, mostly deciduous; inner (lateral) pair often saccate [rarely spurred]. Petals 4, rarely absent, free, alternating with the sepals, usually clawed, arranged in the form of a cross (cruciform, hence the family name), imbricate or contorted [rarely circinate] in bud, entire [rarely lobed, pinnatifid, or fimbriate]. Stamens 6, tetradynamous (the outer 2 shorter than the inner 4), rarely equal in length (Warea) or in 3 pairs of unequal length (Streptanthus), sometimes 2 or 4 (Lepidium) [very rarely 8-24, as in Megacarpaea polyandra), filaments filiform, sometimes winged or appendaged at the base, free [or those of the median pairs of stamens sometimes connate]; anthers 4-sporangiate, 2-loculate at anthesis; pollen grains tricolpate (5- to 7-colpate in Lesquerella) [or up to 10-colpate in Dimorphocarpa], usually oblate or prolate, reticulate, trinucleate when shed. Nectar glands receptacular in origin, highly diversified in shape, size, and disposition around the bases of filaments. Gynoecium 2-carpellate, syncarpous; style persistent, distinct or ob- solete; stigma terminal, capitate or discoid, entire or 2-lobed, the lobes opposite the placenta (replum), rarely opposite the valves, sometimes decurrent and/or connate along entire length; ovary superior, usually sessile (long-stipitate in Warea), 2-loculate, rarely uniloculate, with a false septum connecting the 2 parietal (or rarely subapical) placentae; ovules anatropous or campylotropous, 2-integumented, crassinucellate or tenuinucellate, few to many (sometimes 1) per locule. Fruit typically a bivalvate capsule dehiscing longitudinally from below (commonly called a silique (siliqua) when more than 3 times longer than broad, or a silicle (silicula) when clearly shorter than 3 times the width, but such distinctions arbitrary and sometimes misleading), or fruit indehiscent and becoming lomentaceous or achenelike [or a nutlet, samara, schizocarp, or even a drupe], usually beakless, or having seedless or |- to few-seeded beaks; replum (the framelike placenta) persistent; septum complete (incomplete or reduced to a rim in Armoracia), usually membranaceous [rarely thick and fibrous or 1984] AL-SHEHBAZ, CRUCIFERAE 345 nerved]; gynophores generally lacking or very short (well developed in Warea and Lunaria). Seeds without endosperm, uniseriately or biseriately arranged, winged or wingless, mucilaginous or not when wet; megagametophyte (embryo sac) of the Polygonum type; embryogeny of the Onagrad type; embryo oily, occupying the entire seed, strongly curved (straight only in Leavenworthia) or folded in one of seven ways, most commonly notorrhizal (cotyledons incum- bent—i.e., radicle lying on the back of 1 cotyledon), pleurorrhizal (cotyledons accumbent—radicle applied to the margin of both cotyledons), or orthoplocal (cotyledons conduplicate— folded longitudinally around the radicle) [or diple- colobal—cotyledons twice transversely folded]; germination epigeal. x = 4-13. (Including Raphanaceae Horan., Cruciaceae Dulac.) Type GeNus: Brassica L. A large family of approximately 340 genera and more than 3350 species in some ten poorly defined tribes, distributed throughout the world, primarily in the temperate regions and most successfully in the arid areas (although a few species of Draba and of some other genera have penetrated well into the Arctic and to the subantarctic islands, while others grow at altitudes of up to 6000 meters (19,700 feet) in Kashmir and Tibet). The family is clearly most abundant in the Northern Hemisphere, with the major center of diversification and endemism in the Irano-Turranian region, where some 150 genera (62 endemic) and 900 species (530 endemic) are found, and a secondary center in the Med- iterranean region, with more than 110 genera (21 endemic) and nearly 630 species (290 endemic). Most of the 37 endemic genera and more than 600 species native to North America are distributed primarily in the western United States and northern Mexico. In the Southern Hemisphere, there are 32 endemic genera and some 340 species native to South America (particularly along the Andes and in Patagonia), eight genera and 110 species in South Africa, and 19 genera and 114 species in Australia and New Zealand. Only two genera (Ro- manschulzia O. E. Schulz of Mexico and Central America and Oreophyton O. E. Schulz of eastern tropical Africa) are endemic to the high mountains of the tropics. Of all the genera in the family, only Cardamine L., Lepidium L., and Rorippa Scop. are represented by indigenous species on all continents but Antarctica. Of the 121 species occurring in the southeastern United States, 55 (falling in 27 genera) are naturalized weeds, most of which were originally introduced from Europe. The majority of our 66 native species occur elsewhere in eastern North America, but only 16 of them are endemic to the Southeast, and six others have their centers of distribution in our area. Only one genus (Warea Nutt.) is endemic, and another (Leavenworthia Torrey) has its center of diversity in the Southeast. Seven of the nine taxa of Cakile Miller native to North America occur in our area, but the genus eae as and has diversified in the Old World (southwestern Asia and Eur A very natural family easily distinguished by the eee corolla, the tetradynamous stamens, and the characteristic siliques (hereafter to include the silicle), the Cruciferae— whether placed in the order Capparales of recent au- thors or in the abandoned Englerian Rhoeadales (see Wettstein)— have always been closely associated with the Capparaceae. Systematists are now in agree- ment that the Rhoeadales represent two unrelated orders, the Capparales (con- taining glucosinolates, myrosin cells, and centrifugal stamens, and lacking 346 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 laticifers and benzylisoquinoline alkaloids) and the Papaverales (with benzyl- isoquinoline alkaloids, laticifers, and centripetal stamens, and lacking gluco- sinolates and myrosin cells) (see Gershenzon & Mabry; Rodman, 1981). The family has recently been considered to be a direct descendant from the cap- paraceous subfamily Cleomoideae via the cruciferous tribe Stanleyeae (= The- lypodieae) (Janchen; Takhtajan) or through the Hesperideae (Dvorak, 1973). Although the morphological evidence very strongly favors a connection through the Thelypodieae, none of the extant crucifers is truly archaic, and the paly- nological data (Al-Shehbaz, 1973) do not support such a direct link. It is, therefore, more reasonable to assume that the connection between the two families is through a common ancestor. Fossil evidence is of no help here, and the few scattered reports of cruciferous pollen from the Upper Miocene of France (Muller) and the Cretaceous of New Zealand (Couper), as well as the fossil siliques of Th/aspi L. from the Oligocene of Montana (Becker) and those of Lepidium and other genera (Schulz), only complicate the problem. The tribal classification of the family has occasioned more controversy than any other aspect of its systematics, and none of the existing tribal systems comes close to satisfying all concerned. Although much criticized for its arti- ficiality, Schulz’s is the latest comprehensive monograph on the family, and despite its weaknesses, his system is the most widely followed. With the ex- ception of the tribes Brassiceae, Lepidieae, and Thelypodieae (the Stanleyeae, Streptantheae, and Romanschulzieae of Schulz), all the other large tribes of his system fall short of being natural. Janchen reduced the 19 tribes of Schulz to 15, and his merging of the Matthioleae with the Hesperideae and of both the Drabeae and Lunarieae with the Alysseae is probably justified. The Cremo- lobeae R. Br. (two genera and 36 species of South America) and the Heliophileae DC. (four genera and 77 species of South Africa) are sufficiently distinct and probably merit recognition, but the tribal status of the Pringleeae Hayek and Chamireae Sonder, both unigeneric and monotypic, needs careful evaluation. On the other hand, the Stenopetaleae O. E. Schulz, an Australian unigeneric tribe with eight species, rest solely on the narrowly linear or filiform petals that are circinate in bud, a feature independently evolved in the unrelated Australian monotypic Carinavalva Ising and the North American Lyrocarpa Hooker & Harvey. The genera of the Schizopetaleae R. Br. ex Barn., Schizopetalon Sims (seven species, Chile), Ornithocarpa Rose (two species, Mexico), and Dryope- talon Gray (four species, northwestern Mexico and the adjacent United States), are united mainly by the divided petals. However, they are so different in their fruits, sepal orientation, indumentum, nectaries, stigmas, and cotyledons that they form a highly heterogeneous group of unrelated genera (see Rollins, 1969). In my opinion, each of these genera can be loosely associated with certain members of one of three other tribes. Avetisian (1976, 1983) has recently reduced Schulz’s tribes to three, Thely- podieae, Brassiceae, and Sisymbrieae, but such an action does not seem to be well founded. In the past, tribes have been erected on the basis of differences in a few characters such as cotyledonary position, type of pubescence, and fruit length and the type of its flattening, but this always leads to artificiality in the tribal classification. Perhaps a more realistic classification of the family can be 1984] AL-SHEHBAZ, CRUCIFERAE 347 achieved by grouping the closely related genera and working upward to more natural infrafamilial taxa. The modified version of Janchen’s system adopted here is only intended to provide a workable framework for infrafamilial sub- divisions above the generic level, but even though it represents a major 1m- provement over Schulz’s system, it cannot be considered natural as far as the tribal limits of the Alysseae, Arabideae, Hesperideae, and Sisymbrieae are concerned. Generic boundaries in the family are often arbitrarily drawn, and the establishment of clear-cut intergeneric relationships is often difficult. Al- though there is an average of about ten species to a genus, the majority of genera (250) are oligotypic with five or fewer species, and 138 of these are monotypic. However, more than half of the species of the family belong to 12 large genera: Draba L. (340), Erysimum L. (180), Cardamine (175), Lepidium (175), Alyssum L. (170), Arabis L. (170), Sisymbrium L. (90), Lesquerella S. Watson (80), Rorippa (75), Thiaspi (75), Heliophila L. (72), and Hesperis L. (60). Unlike many of the small genera, the species are generally very distinct throughout the family. A few exceptions, however, do exist, and the most notable examples are the Old World genera Jsatis L., Aethionema R. Br., and Biscutella L., in which hybridization, polyploidy, and apomixis, alone or to- gether, may have played an important role in making species determination a very difficult task. Chromosome numbers have been reported for more than 1400 species (41% of the family total) in 197 genera (author’s compilation). A continuous series of base chromosome numbers from four to 13 exists, but a surprisingly high seen (37%) of the species appear to be based on eight. The lowest chro- ome number known for the family (” = 4) has been found so far only in Ge unrelated genera, the Australian Stenopetalum R. Br. ex DC. and the western North American Physaria (Nutt.) Gray, while the highest number reported (n = 128) is in Cardamine laciniata (Muhl. ex Willd.) Wood. Nearly 37 percent of the species are polyploid, and some of the genera such as Crambe L. (Brassiceae) and Streptanthus Nutt. (Thelypodieae) appear to be exclusively polyploid. Both genera are generally considered the most advanced in their respective tribes. However, Mukherjee believes that aneuploidy and diminu- tion in chromosome size, rather than polyploidy, have played an important role in the evolution of the family. The tribes Alysseae and Arabideae have a base chromosome number of eight, which has been found in more than 60 percent of their species; only about 10-15 percent of their species are based on seven. On the other hand, the tribes Hesperideae, Lepidieae, and Sisym- brieae are based primarily on seven, which has been found in about 40-45 percent of their species, and secondarily on eight, encountered in about 20 percent of the species of each tribe. No single base chromosome number dom- inates in the Brassiceae, and with more than 77 percent of its species known cytologically, the base numbers 7, 8, 9, 10, 11, 12, and 15 occur with frequencies ranging from eight to 20 percent. Although the Thelypodieae have a continuous series of haploid chromosome numbers of ten to 15 (with nearly 46 percent of the species known cytologically), = 14 occurs in more than 60 percent of the species and m = 13 in about 20 percent. Many genera of the Cruciferae have been studied for their chemical con- 348 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 stituents, especially the glucosinolates (mustard-oil glucosides) and the seed fatty acids, both of which have been thoroughly surveyed in the economic species and their wild allies. The fatty-acid composition is known for at least 165 species in 70 genera; in this small sample, the linolenic or erucic acids are the most dominant constituents in the seeds of about 85 percent of the species surveyed. All Cruciferae appear to have glucosinolates, and of the approxi- mately 85 types known, only methyl! glucosinolate (typically characteristic of the Capparaceae) has not been found in any crucifer (see Hedge ef a/.). The distribution of these compounds has been shown to be a valuable tool in chemosystematic studies at the generic and specific levels (Rodman, 1981). In all, some 350 species in about 70 genera have been surveyed, but since most of the earlier reports have dealt only with the distribution of the major con- stituents, many of the species need to be reinvestigated in order to have a complete profile of their glucosinolates. It is agreed that the glucosinolates probably play the most important role in the chemical defense of crucifers against pathogens and herbivores. Research on the distribution of the fatty acids or the glucosinolates has so far failed to provide any meaningful support for the tribal classification of the family. The flavonoid chemistry in the family has not received the attention it deserves, and some of the recent works (Bacon) show that such compounds can be equally valuable in systematic studies in this family. Other secondary metabolites generally occur in negligible amounts, and they are often overlooked. However, relatively high concentrations of alkaloids (Lunaria L.), cucurbitacins (/beris L.), and cardenolides (Erysimum) may be found, and the distribution of the last group of compounds may prove to be useful in solving some of the problems in taxonomically difficult genera such as Erysimum. Species of a few genera are known to accumulate high amounts of selenium (Stan/eya Nutt.) or nickel (Alyssum, Streptanthus), but these capacities have no taxonomic value. Floral anatomy in the Cruciferae has been studied in detail, mainly to resolve several controversial aspects of the gynoecial structure, such as the vasculature of the ovary, the number of carpels, the origin of the septum, the position of the stigmatic lobes, and the derivation of the placentae. The widely accepted bicarpellary hypothesis advocated by Arber, Zohary (1948b), Puri (1951), and Alexander states that the cruciferous gynoecium originated through the con- nation of the margins of two lateral carpels to form two median parietal pla- centae, each of which produces an outgrowth that fuses with the other in the center to form the false septum. This hypothesis, however, fails to provide an adequate explanation for the inverted position of the inner vascular bundles of the replum and for the frequent commissural position of the stigma lobes. The tetracarpellary views of Saunders, Eames & Wilson, Puri (1941), Merx- miiller & Leins, and (more recently) Eigner agree that the cruciferous gynoecium is composed of four carpels, of which the median two are fertile and the lateral two are sterile. Despite the fact that these hypotheses provide sound interpre- tation for the vasculature of the stigma and replum, they do not adequately explain the origin of the false septum and the position of the ovules. Further- more, as indicated by Zohary (1948b), in crucifers with dehiscent fruits, the lines of dehiscence appear only in the later stages of development and do not 1984] AL-SHEHBAZ, CRUCIFERAE 349 correspond with carpel margins, as proposed by Saunders and others (see above). Although I support the bicarpellary interpretation and believe that the gynoe- cium of the Cruciferae is homologous to that of the closely related Capparaceae, both the bi- and tetracarpellary views fail to provide fully satisfactory answers for all the various aspects of the cruciferous gynoecium. The reader is advised to consult Maule and Roth for further details. Flower colors of Cruciferae are predominantly yellow, white, and shades from lavender to purple; true blue or red flowers, if they occur at all, are indeed very rare. The remarkable constancy of floral architecture in the family has been very closely linked to pollination by insects, particularly various Hy- menoptera, Diptera, Lepidoptera, and some Coleoptera. Wind pollination 1s extremely rare and is probably best known in Pringlea antiscorbutica R. Br., a species restricted to the small Kerguelen and Crozet islands of the southern Indian Ocean. Autogamy is common in many of the weedy species, while cleistogamy always occurs in the submersed plants of Subularia aquatica L. Unlike protogyny, protandry appears to be rare in the family (Al-Shehbaz, 1977), and except for very few examples, the flowers of the Cruciferae are almost always perfect. Dioecism is known in three species of Lepidium from New Zealand, while monoecism has been reported in Megacarpaea megalo- carpa (Fischer ex DC.) Schischkin ex Fedtsch., of central Asia and southeastern Russia. Fruits of the Cruciferae are so diverse that they are the most reliably used structures for the proper identification of genera and species. In plants with dehiscent siliques, seed dispersal—even if explosive, as in Cardamine—is con- fined to short distances from the parent plant. However, because of their small size, seeds of the family in general are easily washed farther away by rain or transported by strong winds in open habitats. The corky fruits of all but one of the taxa of Cakile and of some species of Crambe and Raphanus L. are transported by sea, while the winged seeds and samaroid and bladdery fruits that have independently evolved many times in the family are dispersed by wind. The dustlike seeds of certain Saharan species of Diplotaxis DC. ma weigh as little as 0.05 mg and can therefore be transported by storms for several hundred miles. The remarkable rose of Jericho (tumbling or resurrection mus- tard), Anastatica hierochuntica L., 1s dispersed by the tumbling of the entire dry plant, and this species has a continuous distribution extending 8000 ki- lometers (5000 miles) in hot deserts from Mauretania to western Pakistan. Fruit dispersal by mammals is known for several genera having hooked hairs (Tauscheria Fischer ex DC.), glochidiate spines (C/ypeo/a L.), and other adap- tive features. Geocarpy has evolved independently in the Australian Geococcus pusillus Drumm. ex Harvey, in the South American Cardamine chenopodiifolia Pers., and in Morisia monanthos (Viv.) Ascherson, of Corsica and Sardinia. The family is primarily herbaceous, and only some five percent of its species are typically woody; more than 62 percent are perennials. Growth forms, how- ever, may vary from delicate annual herbs to the South American hummock- forming Xerodraba pycnophylloides (Spegaz.) Skottsb. and Lithodraba men- dociensis (Spegaz.) Boelcke, the South African woody climber Heliophila scandens Harvey, the large shrubs Fo/eyola Maire of northern Africa and Par- 350 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 olinia Webb of the Canary Islands, or even the small tree Farsetia somalensis (Pax) Gilg & C. Benedict, of Somalia, Ethiopia, and Kenya. Farsetia Turra also contains a few annual or perennial herbs, which may be less than 10 cm high, as well as large shrubs that may exceed 2 m in height in eastern tropical Africa. Typical shrubs are found in about two percent of the total species of Cruciferae in at least 16 genera scattered in different tribes. The woody con- dition appears to be almost always a derived one, and it must have evolved independently many times within the famil The family includes a number of important crop plants that are grown as food for humans or animals, as sources of condiments or edible and industrial oils, or as ornamentals. The family is also known for its more than 120 weedy species of local or cosmopolitan distribution that invade cultivated lands and occupy disturbed sites, roadsides, waste grounds, and the like. Brassica is the most important genus, for it contains several vegetable and salad plants such as cabbage, cauliflower, Brussels sprouts, kale, broccoli, kohlrabi (all considered to be varieties of B. oleracea L.), turnip and Chinese cabbage (B. campestris L.), rape (B. napus L.), Chinese or Indian mustard (B. juncea (L.) Czern.), and black mustard (B. nigra (L.) W. D. Koch). Other crops include the radish (Raphanus sativus L.), watercress (Nasturtium officinale R. Br.), and the com- mon or garden cress (Lepidium sativum L.). Condiments are obtained from the fleshy roots of horseradish (4rmoracia rusticana Gaertner, Meyer, & Scherb.), while table mustard is prepared from a mixture of the seeds of the white mustard (Sinapis alba L.) and those of either the black or the Indian mustard. Oils from crucifer seeds, particularly from rape, rank fifth in terms of the world tonnage production, and most of it (except that used for making mar- garine in Europe and cooking oil in India) is utilized in the manufacture of numerous industrial products. The seed cake remaining after the expression of oil is rich in protein, and until recently it has been extensively used as feed for farm animals. It contains potentially harmful mustard oils, however, so this usage has become very limited, and most of the seed cake 1s now used as a fertilizer. The most important ornamental crucifers include the wallflower (Erysimum Cheiri (L.) Crantz, rocket or dame’s violet (Hesperis matronalis L.), candytuft (Iberis species), honesty or money plant (Lunaria annua L. and L. rediviva L.), sweet alyssum (Lobularia maritima (L.) Desv.), stock (Matthiola incana (L.) R. Br.), aubrietia (Aubrieta deltoidea (L.) DC.), rock cress (Arabis species), and some species of the genera Aethionema, Alyssum, Brassica, and Draba. The historic blue dye woad was obtained from the fermented ground leaves of /satis tinctoria L. REFERENCES: The volume of literature dealing with the various aspects of the family 1s enormous. Most of the important literature published before 1936 is thoroughly covered by Schulz and has not been repeated here. Although the number of references given here may seem excessive, only about half of the literature consulted during the preparation of this paper has been included. 1984] AL-SHEHBAZ, CRUCIFERAE 351 ALEXANDER, I. Entwicklungsstudien an Bliiten von Cruciferen und Papaveraceen. Planta 41: 125-144. 1952. [Floral development in Arabis, Cheiranthus, and Hypecoum;, “homology” in floral diagram between Cruciferae and Fumariaceae AL-SHEHBAZ, I. A. The biosystematics of the genus Thelypodium (Cruciferae). Contr. Gray Herb. 204: 3-148. 1973. [Generic limits, keys, and evolutionary trends among members of the tribe Thelypodieae; pollen of selected Capparaceae; distribution of mustard oils.] ——. Protogyny in the Cruciferae. Syst. Bot. 2: 327-333. 1977. [Report of protogyny in 21 species of 19 genera, and a compilation of that previously reported in 61 species of 32 genera.] M. M. At-Omar. In: A. LOVE IOPB chromosome number reports LX XVI. Taxon 31: 574-598. 1982. eae 587-589, counts for 58 species in 38 genera. | AppeLovist, L.-A. Lipids in Cruciferae: VIII. The fatty acid composition of seeds of some wild or partially domesticated species. Jour. Am. Oil Chem. Soc. 48: 740- 744. 1971. [Fatty-acid composition of 29 species.] — . Lipids in the Cruciferae. Pp. 221-277 in J. G. VAUGHAN et al., eds., The biology and chemistry of the Cruciferae. London, New York, & San Francisco. 1976. [A review of fatty-acid composition, sterols, waxes, and other lipids; special emphasis on the cultivated species; /9 tables.] Arser, A. Studies in floral morphology. I. On some structural features of the cruciferous flower. New Phytol. 30: 11-41. 193la. [The vascular supply of various floral parts; prefers the bicarpellary origin of gynoecium for descriptive purposes but questions the justification for treating the bi- or tetracarpellary hypotheses as true. dies in floral morphology. II. On n some normal and abnormal crucifers: with Capsella, Crambe, Diplotaxis, Nasturtium, ana Sinapis, strong criticism of Saun- ders’s assumptions that abnormalities represent reversionary ancestral conditions; see SAUNDERS.] ARYAVAND, A. Contribution a l’étude cytotaxinomique de quelques Cruciféres de ’Iran et de la Turquie. Bull. Soc. Neuchateloise Sci. Nat. 98: 43-58. 1975. [Counts for 33 species in 18 genera.] AvETISIAN, V. E. Some modifications of the system of the family Brassicaceae. (In Russian; English summary.) Bot. Zhur. 61: 1198-1203. 1976. [The six largest tribes reduced to two, Brassiceae and Sisymbrieae, the latter encompassing the Alysseae, Arabideae, Hesperideae, and Lepidieae; parallelism in most characters of these tribes : Some peculiarities in distribution of Brassicaceae in connection with their evo- lution. (In Russian.) /bid. 65: 825-829. 1980. [Lists the ee of some 170 genera of the oat a. the highly artificial S n the North Temperate regions of the Old W J . The system of ne cals Brassicaceae. (In Russian; English summary.) /bid. 68: 1297-1305. 1983. [Three tribes recognized: Thelypodieae (including Pringleeae and Cremolobeae), Brassiceae (including Chamireae and Heliophileae), and Si- symbrieae (including the remainder of the family).] Bacon, J. D. Taxonomy of Nerisyrenia (Cruciferae). Rhodora 80: 159-227. 1978. [Ex- tensive account of the cytology and flavonoid chemistry in the genus; nine species recognized. Baez Mayor, A. Estudio cariol6gico . ars cruciferas y su interpretacién en la ee Cavanillesia 6: 59-103. 4. [Basic number for family x = 7; counts r Sisymbrium, Alliaria, ee aes Eruca, Brassica, Moricandia, pherntis Camelina. BaILLon, H. Cruciferae. Hist. Pl. 3: 181-292. 1871. [English translation by M. M. Hartoc, 3: 179-291. 1874.] 352 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 BATEMAN, A. J. Self-incompatibility systems in angiosperms. II]. Cruciferae. Heredity : 53-68. 1955a. [A survey of 182 — s in 80 genera, with particular emphasis on Brassica, Capsella, and Cardami Note on dioecy i in the Cnet. “Tid. 415. 1955b. [Lepidium sisymbrioides 2 e.] Bayer, A. Beitrége zur systematischen Gliederung der Cruciferen. Bot. Centralbl. 18: 119-180. 1905. [Recognizes 15 tribes based primarily on differences in nectar-gland morphology.] Becker, H. F. Oligocene plants from the upper Ruby River Basin, southwestern Mon- tana. Geol. Soc. Am. Mem. 82: I- ee 1961. [Thlaspi primaevum, n. sp., 69, pi. 20, figs. 9-11; excellent imprints of fru BEHNKE, H. D. Sie eve-element characters. cere Jour. Bot. 1: 381-400. 1981. [Sieve- tube plastids 1 in Brassicaceae of the S-type in 22 species of 21 genera, and of the P-type in two species of one genus.] G. EscHLBeck. Dilated cisternae in Capparales—an attempt towards the char- acterization of a specific endoplasmic reticulum. Protoplasma 97: 351-363. 1978. BENTHAM, G., & J. D. Hooker. Cruciferae. Gen. Pl. 1: 57-102. 1865. [Treatment by ER. | BERGGREN, G. Atlas of seeds and small fruits of northwest-European plant species with morphological descriptions. Part 3. 260 pp. Stockholm. 1981. [Cruciferae, 108-145, pls. 70-105; keys based on seed characters for 36 genera and 93 species; descriptions of fruits and seeds for aad and species. BERKUTENKO, A. N., & N. N. GuRzENKov. Chromosome numbers and distribution of rate in the south of the Magadan region. I. (In Russian.) Bot. Zhur. 61: 1595- 1603. 1976. [Alyssum, Arabidopsis, Arabis, Cochlearia, Draba, Erysimum, Les- pied counts for ten species; /0 maps.] BOLKHOVSKIKH, Z., V. Grir, T. MATVEJEVA, & O. ZAKHAREVA. Chromosome numbers . Feporov, ed. (Russian and English prefaces.) 926 pp. Leningrad. 1969. (Brassicaceae, 162-179; chromosome counts through 1964.] Bouman, F. Integument initiation and testa development in some sree Bot. Jour. Linn. Soc. 70: 213-229. 1975. [Brassica, Capsella, Lunaria, Sinapi BRewBAKER, J. L. The distribution and phylogenetic significance of a ieleate and trinucleate pollen grains in the angiosperms. Am. Jour. Bot. 54: 1069-1083. 1967. [Cruciferae, 1078; pollen trinucleate when shed, 16 genera sampled.] Britton, N. L., & A. Brown. An illustrated flora of the northern United States, Canada and the British possessions. Vol. 2.1v + 735 pp. New York. 1897. [Cruciferae, 108- 154; ed. 2, vol. 2, 146-196. 1913.] BUNNING, E. Uber die Differenzierungsvorange i in der Cruciferenwurzel. Planta 39: 126- iS ° “Oo o ee 5 ge oO. 3) 5 Buscu, N. A., ed. Cruciferae. In: V. L. Komarov, ed., Fl. USSR 8: 14-606. 1939. {English translation by R. Lavoorrt, 8: 13-453, 471 ~490. Jerusalem. 1970.] Carus, J. F. The medicinal and evconc crucifers of India. Jour. Bombay Nat. Hist. Soc. 40: 693-712. 1939. [Keys, common names, and use of 40 species in 22 genera.] CALESTANI, V. has pee meee ‘delle Crocifere Italiane. Nuovo Giorn. Bot. Ital. 15: 355-390. Cerri e classificazione delle Crocifere. /bid. 24: 245-290. 1917. [Evolution in the flowers, fruits, and embryos; recognizes 16 subtribes and 14 tribes in the three a A. P. pe. Cruciferae. Syst. Nat. 2: 139- 700. 1821. [95 genera in 21 tribes n five subfamilies (subordines) distinguished on the bases of fruit type and of a eee position in relation to the radicle Cruciferae. Prodr. 1: 131-236. 1824. CaRLQuistT, S.. Wood anatomy of Macaronesian and other Brassicaceae. Aliso 7: 365— 1984] AL-SHEHBAZ, CRUCIFERAE 353 384. 1971. [Cheiranthus (= Erysimum), Crambe, Descurainia, Lepidium, Matthiola, Parolinia, Sinapidendron, Stanleya. Coie, R. A. 1-cyanoepithioalkanes: major products of eee tee eee hydrolysis in certain Cruciferae. Phytochemistry 14: 2293, 2294. 1975. [17 species in 11 genera.] . Isothiocyanates, nitriles, and thiocyanates as products of ei of glucosi- nolates in Cruciferae. /bid. 15: 759-762. 1976. [Quantitative determination of 22 glucosinolate products from seedlings of 74 species in 35 genera. Corsett, G. A. New records for the West Virginia Canine Castanea 38: 214-229. 1973. [61 species in 29 genera of Cruciferae in the state; eight new records; 20 maps.] Couper, R. A. New Zealand Mesozoic and Cainozoic plant microfossils. New Zealand Geol. Surv. Paleont. Bull. 32: 1-88. 1960. [Cruciferous pollen, 47, pl. 7, figs. 1, 2; from Haumurian irae or ?Piripauan (Upper Senonian), found commonly throughout Cretaceous and Tert Crisp, P. Trends in the breeding ee cultivation of cruciferous crops. Pp. 69-118 in J. . VAUGHAN et al., eds., The biology and chemistry of the Cruciferae. London, New York, & San Francisco. 1976. [Economic importance, breeding systems, polyploidy, future trends; special emphasis on Brassica.] Cronaguist, A. An integrated system of classification of flowering plants. Frontisp. + xviii + 1262 pp. New York. 19 die eee 436-451; Brassicaceae, 446-449; suggests that the connection between the Brassicaceae and Capparaceae might be found among the Old World ae oki of family occur in Oligocene and more recent deposits. ] Crove._o, T. J., & D. C. MILLER. Floristic similarities among 51 regions of the Soviet Union based on the Brassicaceae. Taxon 31: 451-461. 1982. [Numerical study using principal components analysis of distributional data for 736 species of nine tribes.] Curtis, P. J., & P. M. MeApe. Cucurbitacins from the Cruciferae. Phytochemistry 10: 3081-3083. 1971. [Compounds rare in Cruciferae, found in several /beris species and in Lepidium sativum DANIELAK, R., & B. BoRKOWSKI. Biologically active compounds in seeds of crucifers. Part III. Chromatographical search for glucosynolates. Diss. Pharm. coe 21: 563-575. 1969. [Spot tests and R, values for glucosinolates in 155 sp Das, V.S. R., & K. N. Rao. Phytochemical phylogeny of the se Stonit eae from the Capparidaceae. Naturwissenschaften 62: 577, 578. 1975. [Nine genera of Cruciferae and five of Capparaceae tested for 15 phenolic acids; data support the derivation of the former family from the latter.] Davis, G. L. Systematic embryology of the angiosperms. x + 528 pp. New York. 1966. ] DAXENBICHLER, M. E., C. H. VAN ETTEN, F. S. Brown, & Q. Jones. Oxazolidinethiones and volatile isothiocyanates in enzyme- ee seed meals from 65 species of Cru- ciferae. Agr. OO d Chem. 12: 127-130. DvoRAk, F. On relationship in ie family Brassicaceae. Feddes Repert. 82: 357- 372. 1971. [Relationships between Macropodium, Lunaria, Christolea, Er- mania, and Vvedenskyella based on types of cellular pattern of the septum and on gynophore length; table 1 compares floral and fruit characters. ] The importance of the indumentum for the investigation of evolutional rela- tionship in the family Brassicaceae. Slant Bot. Zeitschr. 121: 155-164. 1973. [Mul- ticellular glandular hairs occur in the Hesperideae and the ancestral Cleomoideae of Capparaceae, suggesting a possible link unicellular glandular hairs only in Des curainia. | Eames, A. J., & C. L. Witson. Carpel morphology in the Cruciferae. Am. Jour. Bot. 15: 251-270. 1928. [Postulate that the cruciferous gynoecium evolved from four carpels (two outer ones that are sterile and two inner that are fertile but without locules and their ovules are placed in the locules of the sterile carpels); the septum represents an expansion of the ventral margins of solid carpels.] 354 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Crucifer carpels. /bid. 17: 638-656. 1930. [Further arguments for their tetracarpelary hypothesis; vascular supply of the ovules is derived from inversely ted ventral bundles. Sane N. W. Chromosome numbers of some wera Ohio Cruciferae. Cas- tanea 39-42. 1963. [Counts for 21 species in 18 genera.] ribution patterns of Ohio Seen Ibid. 29: 164-173, 1964. [84 county distribution maps for 78 species in 35 genera.] n illustrated guide to the Cruciferae o Ohio. /bid. 30: 177-191. 1965. [Keys to genera and to species; 47 colored photos of plants; photographs of seeds. ] Eccers, O. Uber die morphologische Bedeutung des Leitbundelverlaufes in den Bliiten der Rhoeadalen und iiber das Diagramm der Cruciferen und Capparidaceen. Planta 24: 14-58. 1935. EIGNER, J. Zur Stempel- und F uciferae) unter neueren Gesichtspunkten der Bliitenmorphologie und der ree: (English summary.) Beitr. Biol. Pflanzen 49: 359-427. 1973 [1974]. [Gynoecium and fruit development of 34 species in 32 genera; support for the tetracarpellary hypothesis of gynoecium; trends in fruit evolution; modification of Janchen’s system; see JAN- CHEN. ERDTMAN, G. Pollen morphology and plant taxonomy. Angiosperms. 539 pp. Stock- holm. 1952. (Corrected reprint with addendum [pp. 541-553]. New York. 1971.) [Cruciferae, 133, 134; concludes from survey of 80 species in 55 genera that the family is + stenopalynous, resembling the Capparaceae but differing from the Fu- mariaceae; see also AL-SHEHBAZ, 1973, and RoLLINs & BANERJEE, 1979.] ErrLincer, M.G., & A. KJAER. Sulfur compounds in plants. Pp. 59-144 in T. J. Masry, _ ALSTON, & V. C. RUNECKLES, eds., Recent advances in phytochemistry. Vol. 1. “New York. 1968. Feeny, P. Defensive ecology of the Cruciferae. Ann. Missouri Bot. Gard. 64: 221-234. 1977. [Role of glucosinolates in defending crucifers against bacteria, fungi, insects, and mammals; a as attractants to adapted enemies. FERNALD, M. L. Gray’s manual of botany. ed. 8. Ixiv + 1632 pp. New York. 1950. [Cruciferae, 685-728. GERSHENZON, J., & T. J. MAsry. Secondary metabolites and the higher classification of angiosperms. Nordic Jour. Bot. 3: 5-34. 1983. [Glucosinolates in Brassicaceae and other families, 14-16; rable 1 compares the systems of Cronquist, Takhtajan, Thorne, and Dahlgren with respect to the glucosinolate-containing families.] Giic, E., & R. MuUScCHLER. Aufzahlung aller zur Zeit bekannten siidamerikanischen Cruciferen. Bot. Jahrb. 42: 437-487. 1909. [Enumeration of 287 ee 42 genera, four new genera, and 19 new species; numerous new combination GOERING, K. J., R. Estick, & D. L. BRELSForD. A search for high erucic * acid containing oils in the Cruciferae. Econ. Bot. 19: 251-256. 1965. [Seeds of a species in 20 genera tested for percentages of erucic acid, C,, acids, oil, and protein.] GomeEz-Campo, C. Studies on Cruciferae IV. Chorological notes. Anal. ‘Tn st. Bot. Ca- vanilles 34: 485-496. 1978. [Notes on 32 taxa of 18 genera; traces the origin of two species of Brassica adventive to Australia and California. ; & L. DELGADo. Radioresistance in crucifers. Rad. Bot. 4: 479-483. 1964. [More than half of 47 species in 38 genera showed high radioresistance; nuclear volume and chromosome size assumed as significant factors accounting for resistance. ] GUNTHART, A. Beitrage zur Bliithenbiologie der Cruciferen, Crassulaceen und der Gat- tung Saxifraga. Bibliot. Bot. 11(Heft 58). ix + 97 pp., 7/ pls. 1902. aie 3- 38, pls. 1-5; detailed descriptions of the anthesis of 51 species in 14 gen . Uber die Entwicklung und Entwicklungsmechanik der Grintecabiute a ihre Function unter natiirlichen und kiinstlichen Bedingungen. Beih. Bot. Centralbl. 35(Heft 1): 60-170. 1917 ] nhl nQ : ( Cr 1984] AL-SHEHBAZ, CRUCIFERAE 355 Guyot, M. Virescence et organisation de la fleur des Cruciféres. Bull. Soc. Bot. France 109: 170-176. 1962. [Theoretical floral diagram based on teratology.] HaAnnic, E. Untersuchungen tiber die Scheidwande der ae eae Bot. Zeit. 59: 207-245. pls. 8-10. 1901. [Septum as an outgrowth fro rpel margin, appearing at maturity of fruit as two epidermal layers firmly eee at separating at regions of attachment to replum, where they carry stomata; the space between the two layers is filled with loose parenchyma; numerous examples. HARTWELL, J. L. Plants used against cancer. A survey. Lloydia 32: 79-107. 1969. [Cru- ciferae, 79-92; some 40 species of 27 genera; a table of species, common names, medical preparations, disease conditions, comments, and old literature.] — X., A. ss MacLeop, & M. Moreau. Glucosinolates of nine Cruciferae and Ze) Cappar ceae species. Phytochemistry 20: 2355-2358. 1981. [Alyssum, Arabis, OMe Digi, Lobularia, Sinapis, Sisymbrium, Cleome, Gynandropsis.] Hayek, A. von. Entwurf eines es auf phylogenetischer Grundlage. Beih. Bot. Centralbl. 27: 127-335. p/s. 8-12. 1911. [Recognizes ten tribes and 28 subtribes largely circumscribed on characteristics of myrosin cells and nectar glands; descriptions for 231 genera; phylogeny within tribes Hese._, M. The mustards and related plant families in eastern Tennessee. Jour. Tenn. Acad. Sci. 8: 332-340, 382-417. 1933. [Phylogeny and taxonomy of Papaveraceae, Fumariaceae, Cruciferae, and Capparaceae; Cruciferae, 388—406.] Hepce, I.C. As joa ao geographical survey of the Old World Cruciferae. Pp. 1-45 in J. G. VAUG al., eds., The biology and chemistry of the Cruciferae. London, New York, % ne Francisco. 1976. [Evalvaron of me endemism and distributional patterns in Old World regions , and oe trends; 6 tables, 8 figures; tribal dassiiieation: accor dine to five system , A. KJAER, &O. Matver. Dipterygium—Cruciferae or ne ae Notes Bot. Gard. Edinburgh 38: 247-250. 1980. [Presence of methyl glucosinolate in D. glau- cum supports the capparaceous disposition of the es : & K. H. ReEcHINGER. Cruciferae. Jn: K. H. RECHINGER, ed., Fl. Iranica 57: 1- 372. pls. 1-36. 1968. [In Latin; area covers Iran, epee and portions of Pakistan, Iraq, and USSR.] HEGNAUER, R. Chemotaxonomie der Pflanzen. Vol. 3. 743 pp. Stuttgart. 1964. [Cru- ciferae, 586-607, 663-665, 674, 675.] Hewson, H. J. Brassicaceae. Jn: B. G. Bricas et al., eds., Fl. Australia 8: 231-357. 1982. Heywoonp, V. H., ed. Cruciferae. Jn: T. G. Tutin et al., eds., Fl. Europaea 1: 260-346. 1964. HILDEBRAND, F. Vergleichende Untersuchungen tiber die Saftdriisen der Cruciferen. Hinata, K., & T. Nisuio. Self-incompatibility in crucifers. Pp. 223-234 in S. TSUNODA et al., eds., Brassica crops and wild allies. Tokyo. 1980. [Special emphasis on cul- tivated mieniber Horovitz, as &Y. CoHEN, Ultraviolet refl flowers of crucifers. ur. Bot. 59: 706-713. 1972. a patterns of 28 species in “34 genera; 2/ figures ; INampaR, J. A., & N. V. Rao. Light and scanning electron microscopic studies on trichomes of some Brassicaceae. Feddes Repert. 94: 183-190. 1983. [32 species in 20 genera; | SEM plate.) INGRAM, D. S., B. A. KniGuts, I. J. M y, & P. McKay. Studies in the Cruciferae. Changes in ae composition of ne es a faction following germination. Phyto- chemistry 7: 1241-1245. 1968. [Brassica, Cheiranthus, Raphanus, Sinapis.] IvERSEN, T.-H. The morphology, occurrence, and distribution of dilated cisternae of the endoplasmic reticulum in tissues of plants of the Cruciferae. Protoplasma 71: 467- 396 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 477. 1970. [Dilated cisternae found in 35 species of Cruciferae but not in Resedaceae (four spp.) or Papaveraceae (20 spp.).] . BAGGERUD. Myrosinase activity 1 of Brassicaceae Zeitschr. ee 97: 399-407. 1980. rane cultures of seven specie JAFRI, S. M. H. hase In: E. Nasir & S. I. Aut, eds., Fl. West Pakistan 55: 1- pee : 4 1 JANCHEN, E. Das System der Cruciferen. Osterr. Bot Zeitschr. 91: 1-28. 1942. [Criticism and modification of Schulz’s system; 15 tribes and 41 subtribes recognized; see SCHULZ. ] JARETZKY, R. Untersuchungen iiber Chromosomen und Phylogenie bei einigen Cruci- feren. Jahrb. Wiss. Bot. 68: 1-45. 1928. ziehungen zwischen Chromosomenzahl und Systematik bei den Cruciferen. Ibid. 76: 485-527. 1932. [Chromosome counts for 44 Old World genera.] JART, A. The fatty acid composition of various cruciferous seeds. Jour. Am. Oil Chem. Soc. 55: 873-875. 1978. [27 species of Cardamine, Draba, Erysimum, Malcolmia, and Matthiola.] Jones, S. B., Jk. Mississippi flora. IJ. Distribution and identification of the Brassicaceae. Castanea 40: 238-252. 1975. [Keys and distribution maps for 39 species in 24 genera.] sere an Some tropical African Sens Chromosome numbers and taxonomic nts. Bot. Not. 129: 123-130. ; : Gries In: R. M. POLuicy, a Fl. Trop. East Africa. 73 pp. Rotterdam. 1982. [Area covers Kenya, Tanzania, and Uganda; 21 genera and 53 species.] JORGENSEN, L. B. Myrosin cells and dilated cisternae of the endoplasmic reticulum in the order Capparales. Nordic Jour. Bot. 1: 433-445, 1981. [The ultrastructure of protein- -rich cells, idioblastic myrosin cells, or cells of similar appearance supports families Capparaceae, Brassicaceae, Tovariaceae, Resedaceae, and Moringaceae; 70 species investigated. ] KERBER, E. von, & G. BUCHLOH. Sinapinfreie und sinapinhaltige Brassicaceae (= Cru- ciferen). Beitr. Biol. Pflanzen 55: 377-384. 1981. [Seeds of 100 species in 45 genera analyzed for the presence of sinapine; 46 species in 25 genera contain this alkaloid.] Kincssury, J. M. Poisonous plants of the United States and Canada. xiii + 626 p Englewood Cliffs, New Jersey. 1964. [Cruciferae, 28, 29, 158-171; 13 species nee genera. Kyaer, A. Naturally derived isothiocyanates (mustard oils) and their parent glucosides. Fortschr. Chem. Organ. Naturstoffe 18: 122-176. 1960. [Structure and distribution of 30 mustard oils in some 150 species of 47 genera of Cruciferae and in 26 species of ten other familie : Cie ee in the Cruciferae. Pp. 207-219 in J. G. VAUGHAN et al., The biology and chemistry of the Cruciferae. London, New York. & San a 1976. KniGuTs, B. A., & A. M. M. Berrie. Chemosystematics: seed sterols in the Cruciferae. Phytochemistry 10: 131-139. 1971. [Five classes of sterols surveyed in 55 taxa of 21 genera = nine tribes. KNosiocn, I. Intergeneric hybridization in flowering plants. Taxon 21: 97-103. 1972. es 100; 257 interspecific and intergeneric hybrids reported for the family, list of intergeneric hybrids incomplete.] KnutTH, P. Handbook of flower pollination. (English translation by J. R. A. Davis.) Vol. 2. vul + 703 pp. Oxford. 1908. [Cruciferae, 74-128. In German ed., 2(1): 17-130. 1898. Ko sg, K.-P. Serologischer Beitrage zur Systematik der Capparales. (English summary.) Bot. Jahrb. 99: 468-489. 1978. [Capparales and Papaverales show no serological 1984] AL-SHEHBAZ, CRUCIFERAE bey relationships; proposes removal of Moringaceae from the Capparales, the latter consisting of Capparaceae, Brassicaceae, Tovariaceae, and Resedaceae on . Serological investigations to the structure of the Brassicaceae. (In German; English summary.) Pl. Syst. Evol. 140: 39-55. 1982. [21 genera of eight tribes; serological support for tribal aaa of some genera but not others; realignment of the positions of some tri Kron, M. An electron ee study of the behavior of Cruciferae pollen after pollination. Pp. 221-224 in H. F. Linskens, ed., Pollen physiology and fertilization. Amsterdam. 1964. [Brassica nigra.] Kumar, P. R., & S. TsuNopA. Variation in oil content and fatty acid composition among seeds from the Cruciferae. Pp. 235-252 in S. Tsunopa et al., eds., Brassica crops and wild allies. Tokyo. 1980. [A survey of 165 species (not 173 as listed by the authors) in 70 genera.] LaPorte, J. Semillas y plantulas de las Cruciferas cultivadas en la Argentina. 141 pp Buenos Aires. 1959. [Keys, descriptions, and illustrations; species of 24 genera] protein free amino acids in 26 species of Cruciferae and 14 of Resedaceae; methods of analysis, structural determination, biosynthesis, biogenesis; bibliography of 285 references. | Lens, K., & C. HoLzaApFeL. Flora of the Canary Islands: the Cruciferae, the Crassulaceae and the ferns and their allies. Anal. Inst. Nac. Invest. Agrar. Ser. Prod. Veg. 4: 165- 273. 1974. [Cruciferae, 167-201. MacLeop, A. J. Volatile flavour compounds of the Cruciferae. Pp. 307-330 in J. G. VAUGHAN et al., eds., The biology and chemistry of the Cruciferae. London, New York, & San Erencisco: 1976. Maire, R. Cruciferae. Fl. Afrique du Nord 12: 139-403. 1965; 13: 5-365. 1967; 14: 1- 153. 1977. Manton, I. Introduction to the eee ae of the Cruciferae. Ann. Bot. II. 46: 509-556. 1932. [Counts for 250 spp. in 80 genera, relationships sen genera, significance of aneuploidy and Aye in the evolution of the family.] Marais, W. Cruciferae. In: L. E. Copp, B. De Winter, D. J. B. ence & H. B. Rycrort, eds., Fl. So. Africa 13: 1-118. 1970. MarkarafF, F. Cruciferae. Jn: G. Hea, Illus. Fl. Mittel-Europa, ed. 2. 4(1): 73-514. 63. Maut_e, A. F. An anomalous form of Sinapis alba. Bot. Tidsskr. 65: 209-224. 1970. Mepve, R. J. The mycorrhizal status of the Cruciferae. Am. Midl. Nat. 109: 406-408. 1983. [24 species in 15 genera examined; two were shown to be mycorrhizal, bringing the total of mycorrhizal crucifers to eight outs in the genera Brassica, Capsella, Lobularia, Lunaria, Raphanus, and Rorip MERXMULLER, H., & P. Letns. Die ner ner me Merem der Kreuzbliitler und Mohngewachse. (English summary.) Bot. Jahrb. 86: 113-129. 1967. [Morphological support for dividing the Rhoeadales into Papaverales and Capparales; androecium of Cruciferae is diplostemonous and gynoecium is tetramerous.] MetTca.re, C. R., & L. CHALK. Anatomy of the dicotyledons. Vol. 1. Ixiv + 724 pp. Oxford. 1950. Sema 79-87. Miter, R. W., F. R. E E, I. A. Wo.rr, & Q. Jones. Search for new industrial oils. XIII. Oils ‘con 102 oe of Cnet ae Jour. Am. Oil Chem. Soc. 42: 817-821. 1965. , C. H. van Erten, C. McGrew, I. A. Wocrr, & Q. Jones. Amino acid com- BOaHOn of seed meals from forty-one species of ‘Cruciferae. Agr. Food Chem. 10: 426-430. 1962. [Quantitative analysis for 14 protein amino acids.] 358 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 MuENSCHER, W. C. Weeds. ed. 2. xviii + 586 pp. Ithaca & London. 1980. [Cruciferae, 229-261, figs. 44-57.] MUKHERJEE, P. Chromosome study as an aid in tracing the evolution of Cruciferae. Cytologia 40: 727-734. 1975. [Counts for 20 species, karyotype analysis for seven species; proposes that the basic number for the tribes Arabideae and Brassiceae 1s six; aneuploidy and diminution in chromosome size played an important role in evolution within the family.] Mu ter, J. Fossil pollen records of extant angiosperms. Bot. Rev. 47: 1-142. 1981. [Brassicaceae pollen from upper Miocene of France, 40. MUuLLIGAN, G. A. Chromosome numbers of Canadian weeds. I. Canad. Jour. Bot. 35: 779-789. 1957. [Berteroa, oo Capsella, Erucastrum, Erysimum, Lepidium, Neslia, Sisymbrium, Thlasp me nee ae family Cruciferae. I. Ibid. 42: 1509-1519. 1964. [Counts for 24 species in 11 genera. —. Chromosome numbers of the family Cruciferae. II. /bid. 43: 657-668. 1965. [Counts for three species of Braya and ten of Cardamine. —. Chromosome numbers of the family Cruciferae. III. /bid. 44: 309-319. 1966. [Counts for 13 species of Draba and seven of Erysimum, agamospermy 1s reported in E. inconspicuum.] Mur ey, M. R. Seeds of the Cruciferae of northeastern North America. Am. Midl. Nat. 46: 1-81. 1951. [Terminology, key to species based on seed characters, seed de- scriptions, and distributions for 118 species in 49 genera; 94 figures. Nerouitzky, F. Anatomie der Angiospermen-Samen. Handb. Pflanzenanat. II. Arche- gon. 10. vi + 365 pp. 1926. (Cruciferae, 140-143.] PALIWAL, G. S. Ontogeny of stomata in some Cruciferae. Canad. Jour. Bot. 45: 495- 500. 1967. [Subsidiary cells and guard cells originate from same protodermal cell following the syndetocheilic type of ontogeny; 12 species of eight genera Pant, D. D., & P. F. Kipwart. Development of stomata in some Cruciferae. oe Bot. II. 31: 513-521. 1967. [Epidermal structure and ontogeny of stomata in ten taxa of Brassica, Coronopus, Iberis, Lobularia, Nasturtium, and Raphanus, anomalous or rare types of stomata. PatmMan, J. A checklist of Florida Cruciferae. Quart. Jour. Florida Acad. Sci. 25: 192- 200. 1962. [Key, common names, and county distributions of 32 native and nat- uralized taxa; list of 13 species of ornamentals and other cultigens. ] & H. H. I:tis. Preliminary reports on the flora of Wisconsin. No. 44. Cruci- ferae — mustard family. Trans. Wisconsin Acad. Sci. Arts Lett. 50: 17-73. 1961. [68 species, 40 aah from the Old World, in 32 genera; keys, descriptions, distributions, 58 maps.] Perry, L. M. Me dicinal plants of East and Southeast Asia: attributed eg and uses. xii + 620 pp. Cambridge, Massachusetts. 1980. [Cruciferae, 110-113. PoLATSCHEK, A. Chromosome numbers and remarks on systematics and distribution of some Brassicaceae from Europe, northern Africa, Asia, and Australia. (In German; English summary.) Phyton Austria 23: 127-139. 1983. [Counts for 46 taxa of 27 genera; distribution, taxonomic notes, typification of certain taxa. ] Ponzi, R. Presenza di cisterne dilatate del reticolo endoplasmico in alcune specie di Brassicaceae. (English summary.) Delpinoa 20: 105-109. 1978. [Dilated cisternae in parenchyma of vascular bundles of five species of Jberis, Lepidium, Lunaria, PRANTL, K. Cruciferae. Jn: A. ENGLER & K. PRANTL, Nat. Pflanzenfam. HI. 2: 145- 206. 1891. [A system of classification based on pubescence divides the family into four tribes and 20 subtribes; brief descriptions for 208 genera; see SCHULz for Prasap, K. Development and organization of gametophytes in certain species of Cru- 1984] AL-SHEHBAZ, CRUCIFERAE 550 ciferae. Acta Bot. Indica 3: 147-154. 1975. [Brassica, Coronopus, Farsetia, Lepid- ium, Malcolmia. eed coat structure and development in certain species of Cruciferae. New Botanist 3: 95 103. 1976. [Cakile, Capsella, Coronopus, Farsetia, Malcolmia, Thlas- The development and structure of basal body in the ovule and seed of certain species of Cruciferae. Bot. Jahrb. 98: 266-272. 1977. [Brassica, Cakile, Capsella, ee Eruca, Farsetia, Lepidium, Matcolmia, Thlaspi.] rphology and histochemistry of the nucellus and endosperm in certain species of Cnt Ibid. 100: 536-541. 1979. [Endosperm development is nuclear; nu- cellar tissue is persistent and closely associated with haustorial portion of endosperm at chalazal end; histochemical and structural changes; eight genera investigated.] placentation, ‘fertile carpels solid; speculates that the origin of septum 1s receptacular in the basal region and placental in the upper.] . The role of floral anatomy in the ae of morphological problems. Bot. Rev. 17: 471-553. 1951. [Crucifer carpels, 509, 510.] Queiros, M. Contribuicao para 0 conhecimento citotaxonomico das spermatophyta de Portugal. IX. Cruciferae. Bol. Soc. Brot. 47: 315-335. 1973. [Counts for 52 taxa in 30 genera.] Raprorp, A. E., H. E. AHLES, & C. R. BELL. Manual of the vascular flora of the Carolinas. lxi + 1183 pp. Chapel Hill. 1968. [Cruciferae, 486-51 1.] Rao, N. V., & J. A. INAMpDAR. Structure and development of normal and abnormal stomata in the seedlings of some Cruciferae. Proc. Indian Acad. Sci. Pl. Sci. 90: 521-533. 1981. [Anisocytic, anomocytic, helicocytic, paracytic, and anomalous types found in 16 species of nine genera.] Leaf architectural studies in the Brassicaceae. Bot. Mag. Tokyo 96: ne 28. 1983. [Venation types of 35 selected species of 19 genera; tracheids and condary veins show extraordinary variability. ] Rickey H. W. Wild flowers of the United States. The ae states. Vol. 2, part x + 322 pp. New York. 1967. [Cruciferae, 220-239, pls. & Roos JE: eee: methods of analysis and some eee ait prob- ms. Phytochem. Bull. 11: 6-31. 1978. [Isolation, separation, identification, and paeeieriee it 176 ache cited. Divergence, convergence, and parallelism in phytochemical characters: the aa cosinolate- myrosinase system. Pp. 43-79 in D. A. YounG & D. S. SEIGLER, eds. Phytochemistry and angiosperm phylogeny. New York. 1981. [Excellent review dealing with aspects of the chemistry, anatomy, genetics, and ecology of these com- pounds; critical evaluation of the taxonomic positions of the 12 glucosinolate-con- taining families within the systems of Cronquist, Dahlgren, and Thorne with respect to parallelism and convergence ‘HEW. Phytochemical correlates of herbivory in a community of native and naturalized Cruciferae. Biochem. Syst. Ecol. 8: 43-50. 1980. [Plant- specific ED pi.| Ro tuins, R. C. Chromosome numbers of Cruciferae. Contr. ine iam 197: 43-65. 1966. [Counts for 113 species in 29 genera; /0 figures in 3 plate. . A remarkable new crucifer from Mexico. /bid. 198: 3-8. 1969. [Ornithocarpa torulosa, sp. nov.; comments on the tribe eee Weeds of the Cruciferae (Brassicaceae) in rth America. Jour. Arnold Arb. 62: 517-540. 1981. [Common names, habitats, St eceeaiel coe ene bers for 95 introduced species in 48 genera and for 26 native species in 15 genera.] 360 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 & U. BANERJEE. Trichomes in studies of the Cruciferae. Pp. 145-166 in J. G. VAUGHAN et al., eds., The biology and chemistry of the Cruciferae. London, New York, & San Francisco. 1976. [Comparative study of trichomes in 69 species of Lesquerella; 36 SEM figures in 6 plates & Pollens of the Cruciferae. Publ. Bussey Inst. Harvard Univ. 1979; 33- 64. 1979. [14 SEM plates of pollen of 35 species in 26 genera; see also ERDTMAN.] & L. RiipENBERG. Chromosome numbers of ene I. Contr. Gray Herb. 201: 117-133. 1971. [Counts for 59 species in 20 genera; 28 figures in 5 plates.] Chromosome numbers of Cruciferae UL bid 207: 101-116. 1977. [Counts for 57 species in 29 genera; /8 figures in 4 plates.] & romosome numbers of Cruciferae IV. Publ. Bussey Inst. Harvard Univ. 1979: 79- 92. 1979. [Counts for 34 species in 17 genera; /2 EN ae ae Rotn, I. Fruits of angiosperms. Encyclopedia of plant anatomy. Vol. 10, pt. 1 675 pp. Berlin & Stuttgart. 1977. (Cruciferae, 218-225. Sampson, D. R. Intergeneric pollen-stigma incompatibility in the Cruciferae. Canad. our. Genet. Cytol. 4: 38-49. 1962. [Pollen-tube inhibition on stigmas; 18 inter- es pollinations between 13 species of ten genera.] SATYANARAYANA, A., & S. K. SEN. A simple and oe squash technique for somatic chromosomes of crucifers. Ibid. 15: 369-371. a. SAUNDERS, E. R. A reversionary character in the ea (Matthiola incana) and its sig- nificance in regard to the structure and evolution of the gynoecium in the Rhoeadales, the Orchidaceae, and other families. Ann. Bot. 37: 451-482. 1923. [Cruciferous gynoecium typically composed of four carpels, but believes that some genera have as many as 50 carpels; evidence based on abnormally developed fruits. ] na new view of the nature of the median carpels in the Cruciferae. Am. Jour Bot. 16: 122-137. 1929. [Criticism of the tetracarpellary gynoecium as interpreted by EAmMes & WILSON (1928).] ScHutz, O. E. Cruciferae. Jn: A. ENGLER & K. PRANTL, Nat. Pflanzenfam. ed. 2. 17b: 227-658. 1936. [The most comprehensive treatment of the family to date; 35] genera classified in 19 tribes and 30 subtribes with keys, descriptions, and 306 figures; the basic reference for the earlier literature on the family. SCHWEIDLER, J. H. Uber den Grundtypus und die pig re cea der Cruci- feren-Nectarien I. Beih. Bot. Centralbl. 27: 337-390. p/. 13. 1911 SHAw, E. A. Taxonomic revision of some Australian endemic genera of Cruciferae. Trans. Proc. Roy. Soc. S. Pesce 89: 145 253. 1965. [Blennodia, Arabidella, Harm- siodoxa, Scambopus, Drabastrum, Pachymitus, Geococcu Revision of reine is (Cruciferae), Jour. Arnold Arb. 53: 52-75. 1972. [Eight species endemic to Australia with haploid numbers of four or five.] . Revisions of some genera of Cruciferae native to Australia. Contr. Gray Herb. 205: 147-162. 1974. [Phlegmatospermum, Cuphonotus, Microlepidium, Ballanti- nia.]| SHIVANNA, K. R., Y. HeEsLop-HaArrison, & J. sLop-HARRISON. The pollen-stigma interaction: a pollination in the ha ae Aeha Bot. Neerl. 27: 107-119. 1978. [The involvement of the coating on the stigmatic papillae in the incompatibility reaction; Raphanus, Cheiranthus, Sinapis.] SIMMONDS, N. , ed. Evolution of crop plants. xii + 339 pp. London & New York. 1976. [Brassica spp., 45-59; alee 56-59; Raphanus, 60-62; Nasturtium, 62-64, Armoracia, 305, 306; Lepidium, 3 SMALL, J. K. Manual of the eouiheasicn flora. xxii + 1554 pp. New York. 1933. [Brassicaceae, 551-575.] SmituH, E. B. An atlas and annotated list of vascular plants in Arkansas. iv + 592 pp. Fayetteville, Arkansas. 1978. [Cruciferae, 123-136, 515, 528. SmitH, F. H. Some chromosome numbers in the Cruciferae. Am. Jour. Bot. 25: 220, 221. 1938. [Counts for 12 species in eight genera.] 1984] AL-SHEHBAZ, CRUCIFERAE 361 Spratt, E.R. The gynoecium of the family Cruciferae. Jour. Bot. London 70: 308-314. 1932. [On the basis of the anterior-posterior position of the stigma and the large size of the replum bundle that extends to the stigma, hypothesizes that the ovules are attached to the midribs of the carpels.] Stace, C. A., ed. Hybridization and the flora of the British Isles. xi11 + 626 pp. London, New York, & San Francisco. 1975. [Pp. 137-153, Brassica, Diplotaxis, es Cakile, Capsella, Cochlearia, Erophila, Cardamine, Barbarea, Arabis, Roripp TAKHTAJAN, A. L. Outline of the classification of flowering plants (Magnoliophyta). Bot. Rev. 46: 225-359. 1980. [Holds the view originally advanced in earlier versions of his system that the Cruciferae are derived from Capparaceae-Cleomoideae through the tribe Stanleyeae (Thelypodieae); Capparales includes five families placed in three suborders.] THEOBALD, W. L., J. L. KRAHuLIK, & R. C. Rotiins. Trichome description and clas- sification. Pp. 40-53, 190-198 in C. R. Metcatre & L. CHALK, Anatomy of the dicotyledons. ed. 2. Vol. 1. Oxford. 1979. [Numerous examples from Cruciferae. ] Tuorne, R. F. Proposed new realignments in the angiosperms. Nordic Jour. Bot. 3: 85- 117. 1983. [Brassicaceae, 104, placed in the Capparales of the superorder Violiflorae; includes the tae Dipterygium in the Brassicaceae. ] Tsunopa, S., K. TA, & C. GOMEz-Campo, eds. Brassica crops and wild allies. xvill + 354 is Tokyo. 1980. [Nineteen papers dealing with various aspects of the cultivated crucifers and their wild relatives. VALDES BERMEJO, E. Estudios citotaxonomicos en Cruciferas espafiolas. Ciencias Madrid 9: 80-84. 1974. VAUGHAN, J.G. The a ale and staining of sections of cruciferous seed coats. Stain Tech. 35: 229-231. 1960. , A. J. am & B. M. G. Jones, eds. The biology and chemistry of the Cruciferae. xvi + 355 pp. oe New York, & San Francisco. 1976. [Eleven papers dealing with various aspects of the family.] ,J.R. : DENFORD. Seed studies in the Cruciferae. Pp. 119-144 inJ. G. VAuGHAN et al., eds., The biology and chemistry of the Cruciferae. London, New York, & San Francisco. 1976. [Seed structure, chemistry; particular emphasis on Brassica.] J. M. Wuitenouse. Seed structure and the taxonomy of the Cruciferae. Bot. Jour. Linn. Soc. 64: 383-409. 1971. [Seeds of 200 species in 90 genera, 10 figures, I plate, seed-coat eas do not support tribal divisions of the family.] Vieci, L., A. M. Pacnt, G. Corsi, & G. CELA RENzoNI. Embryo suspensor in Cruciferae. I. Morphology and ata (In Italian; English summary.) Giorn. Bot. Ital. 110: 347-357. 1976. [Seven species of Alyssum, Brassica, Capsella, Diplotaxis, Eruca, VILLANI, A. Sulla classificazione delle Crocifere. An om -1 1926. eae for 102 genera; 11 tribes recognized, the traditional Hiasciceac divided into ee YY. F. The forms of heterocarpy in the Brassicaceae Burn. family and the evaluation of their evolutionary significance. (In Russian; English summary.) Bo Zhur. 53: 1428-1439. 1968. [Heterocarpy and heterospermy; special emphasis on members of the Brassiceae.] —. Experience in an ecological-geographical and phylogenetic analysis of hetero- carpy (on the example of Cruciferae.) (In Russian; English summary.) Biul. Moskova Obshch. Isp. Pri. Biol. 75(3): 77-84. 1970. [Occurrence of heterocarpy in 41 genera a derived feature restricted to advanced groups; adaptive significance. ] WALLACE, D. H., & M. E. NASRALLAH. Pollination and serological procedures for iso- lating incompatibility genotypes in the crucifers. Mem. Cornell Univ. Agr. Exper Sta. 406: 1-23. 1968.* WaLpo_e, B. A. Distribution of the Cruciferae in Michigan. Pap. Mich. Acad. Sci. Arts 362 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Lett. 6: 307-349. 1927. [Keys, distributions; 34 genera and 81 species, only 28 species indigenous. We su, S. L., & J. L. REVEAL. Utah flora: Sins aoe (Cruciferae). Great Basin Nat. 37: 279- 365. 1977. [Keys, descriptions, distribution data, new taxa and combina- tions, index; 155 species, including 37 Cais co WETTSTEIN, R. Handbuch der systematischen Botanik. ed. 4. x + 1152 pp. Leipzig & Vienna. 1935. [Rhoeadales, 716-728; Cruciferae, 722-727; see ees 12 tribes.] Wiius, J.C. A dictionary of the flowering plants and ferns. ed. 8. (Revised by H. K. Airy SHAW.) xxii + 1245 + Ixvi pp. Cambridge, England. 1973. iCruciferae, 307- 309 ] YARNELL, S. H. Cytogenetics of the vegetable crops. II. Crucifers. Bot. Rev. 22: 81-166. 1956. [Armoracia, Brassica, Crambe, Eruca, Lepidium, Nasturtium, Raphanus, Si- napis, special emphasis on Brassica and Raphanus, 288 references cited.] YEN, C. Ona new view of carpel morphology in Brassica. (In Chinese; English summary.) Acta Bot. Sinica 8: 271 fa 1959. [The six-carpel hypothesis; normal and abnormal fruits of B. napella Choi ZOHARY, M. Follicular ne in Cruciferae. Lloydia 11: 226-228. 1948a. [Lepta- eum filifolium; fruit dehiscence from separation tissue restricted to the adaxial (posterior) side; such tissue lacking on the abaxial side. Carpological studies in Cruciferae. Palestine Jour. Bot. Jerusalem Ser. 4: 158- 165: 1948b. [Use of terms silique and silicle discarded; seven fruit types recognized — valvoid, folliculoid, nucamentoid, valvo-nucamentoid, valvo-lomentoid, lomen- toid, and schizocarpoid; evolutionary trends in fruit; support for the bicarpellary origin of cruciferous gynoecium. | KEYS TO THE TRIBES AND GENERA OF CRUCIFERAE IN THE SOUTHEASTERN UNITED STATES General characters: Mostly annual, biennial, or allies herbs, rarely shrubs, glabrous or with simple or variously branched unicellular trichomes, rarely with multicellular glan- dular trichomes, leaves exstipulate, usually simple, Mee inflorescences basically ra- cemes or corymbs (flowers rarely solitary), usually ebracteate; flowers hypogynous, mostly acinomorpi perfect; nena, in 2 decussate pairs, erect or spreading, the lateral fees r often saccate at the base; corolla cruciform, the petals 4, usually clawed, rarely absent; ae glands receptacular, Saris or subtending the bases of some or all filaments; androecium of 6 stamens (rarely 2, 4, or more than 6) in 2 whorls, the outer pair usually shorter than the 2 inner pairs (tetradynamous), rarely all equal in length; gynoecium of ited carpels,; ovary superior, often 2-locular by a false septum connecting the 2 parietal placentae; style persistent, distinct or obsolete; stigma entire or 2-lobed; ovules I to many, natropous or campylotropous; fruit basically a capsule (often called a silique), og longitudinally by 2 valves, sometimes indehiscent and modified to a loment, nutlet, mara, or schizocarp; seeds without endosperm, winged or wingless, mucilaginous or a when wet, embryos large, almost always folded or curved in one of several ways Key to the Tribes A. Fruits _ beaked, sometimes transversely jointed and breaking at maturity into or more seed-bearing segments; beak |- or few-seeded, rarely seedless; cotyledons almost ae conduplicate. 2.2.0.2... 0.00 eee Tribe 2. BRASSICEAE. A. Fruits beakless or very ss with a seedless stylelike beak, never jointed; cotyledons accumbent or incum B. Fruits strongly compressed at right angles to the septum; see much narrower Aa WAG OF TAO: o2 isn rngae dane nods tamed ed eee ribe 3. LEPIDIEAE. 1984] AL-SHEHBAZ, CRUCIFERAE 363 B. Fruits terete, angular, inflated, or compressed parallel to the septum; replum equaling width of fruit. C. Fruits less than 3 times longer than broad (except in a few species of Draba); plants mostly with stellate or furcate trichomes, with or without unbranched ONC Sere no nts aarp Pe aes ae eae Tribe 4. ALYSSEAE C. Fruits mostly more than 3 times longer than broad, if less the plants (ours) glabrous; trichomes branched, unbranched, or absent. D. St ns mostly exserted, equal in length, sometimes slightly tetradyna- mous, or in 3 pairs of unequal length; petals mostly crisped or channeled; gynophores (1-)2-30 mm long; trichomes simple or absent. .......... Behind ee ae ee ene eee eee ee yas Tribe 1. THELYPODIEAE. . Stamens included or slightly protruding, tetradynamous; petals usually neither crisped nor channeled; gynophores absent or rarely present, to 2 mm long; trichomes branched, simple, or lacking. E. Sepals erect, al closed at anthesis; multicellular glandular trichomes commonly present. ....................0005 ribe 6. HESPERIDEAE. E. Sepals ae or r ascending, calyx open at anthesis; multicellular iv) F. Cotyledons aecunabene Je eee Ge a eee Tribe 5. ARABIDEAE. F. Cotyledons incumbent. ................. Tribe 7. SISYMBRIEAE. Key to the Genera Several species of Jberis L. (candytuft), Ma/colmia R. Br. (Virginian stock), and Mat- thiola R. Br. (stock) are eulivated in our ar ea, os there are no records that any of them has b alcolmia africana R. Br. and Matthiola longipetala (Vent.) DC. are well- iar weeds in Texas i some of the western states, but neither of them has been found in our area. These three genera are included in the following key, but they will not be dealt with further A. Fruits transversely jointed or lomentaceous, indehiscent, often breaking transversely at maturity into 1- or few-seeded segments. B. Plants with stalked, multicellular, glandular trichomes; fruits not transversely jointed; stigmas with strongly decurrent connate lobes. ....... 37. Chorispora. B. Plants eglandular, glabrous or with simple, unicellular trichomes only; fruits trans- versely jointed; stigmas entire or 2-lobed—if 2-lobed, lobes neither decurrent nor connate C. Lower segment of fruit always seedless; upper segment several seeded, more than 10 times longer than the lower segment. ............... 9. Raphanus. C. Lower segment of fruit usually 1-seeded, rarely 3-seeded or seedless; upper segment |- or rarely 2-seeded, equaling or up to 5 times longer than the lower segment. D. Petals yellow; fruits not corky, less than 1 cm long; cotyledons condupli- cate; hirsute, nonfleshy weeds of waste grounds and cultivated land. opine § shes ni laa 5 ee uy te fia Be eee eulogy den cay hy 0. Rapistrum, D. Petals white, or lavender to purple; fruits corky, often more than 1.5 ¢c long; cotyledons accumbent, rarely incumbent; glabrous, often fleshy ane Ofsandy shores ar beaches... .2..5.4..4s452h¢ariowiadduekes 12. Cakile. A. Fruits neither jointed nor oo dehiscent or rarely indehiscent (Calepina), not breaking transversely into segm E. Early flowers solitary, borne on scapes 3-10 cm long originating from center of a basal rosette; radicles straight or tently curved, much shorter than cotyledons. All flowers borne in terminal or axillary racemes or corymbs; radicles strongly curved, about as long as the accumbent, incumbent, or conduplicate cotyledons. m 364 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 F. Fruits less than 3 times longer ~ broad, broadly oblong or elliptic to globose, orbicular, triangular, or didym G. Fruits strongly Seer ‘right angles to the septum; replum much narrower than width of fruit. H. Flowers zygomorphic; outer petals markedly larger than inner. I. Seed | in each locule; median filaments not appendaged. ....... x Sheet saves aneeg ai esate. ductesay aihacrsa? toes nans anes nes tal tareem nate enna [Iberis.] I. Seeds 2 in each locule; median filaments with a broad, oo. a sre eink eg or ae a se ee 18. Teesdalia. ppendage. Flowers actinomorphic; petals equal in size, sometimes ae or lacking. J. Se K. K. J. Se Ly eds 3 or more, rarely 2 per locule. Fruits obcordate-triangular; basal leaves forming a rosette; plants with furcate and stellate trichomes. ........... 19. Capsella. Fruits orbicular or elliptic; basal leaves not in et plants labrous or = unbranched trichomes only. ...17. Thlaspi. ed | per locu . Fruits pene reticulate or verrucose, often didymous; inflo- rescence axillary; upper cauline leaves 1|- or 2- oo sa eM Els aha acdsee ese eaigee ead udse Gate eee eee 15. Cor onopus. Fruits smooth; inflorescence terminal; upper cauline leaves en- tire, toothed, or pinnately lobe M. Fruits dehiscent, obtuse or cuneate at base, retuse or emar- tee at apex; annuals, biennials, or nonstoloniferous pe- 1 sepaaee ara 4thn ton ee dinars ene a eeee eee 4. Lepidium. M. Fruits meee cordate at base, cuneate at apex; stolon- iferous perennials. ....................05. 16. Cardaria. G. Fruits inflated, globular, or compressed parallel to the septum; replum about as broad as width of fruit. N. Plants glabrous or pubescent with unbranched trichomes only. Fruits coarsely reticulate-rugose, sees indehiscent, 1-seeded; petals unequal; cotyledons conduplicate. ........ 11. Calepina. N. O. F ruits smooth, not ribbed, dehiscent, nee seeded; petals equal; cotyledons accumbent. P. ae) with o Fruits strongly compressed i to the septum; seeds com- pressed, broadly winged, 3-10 mm wide. Q. Flowers purple, rarely white; Cee ebracteate; leaves dentate; fruits more than 1.5 cm broad; gynophores 1-3 er 7 ; pinnatisect; fruits less than 1 cm broad; gynophores less than 0.5 cm long; funiculi free from the septum. ....... . ae inflated; seeds plump, wingless, less than 1.5 mm wide. . Flowers white; septum rudimentary or incomplete; plants anes heats ct ie ted nemo 4) de aca paye aok 36. Armoracia. Il pt lete; pl lor biennial, ee erenn nial. re Ce a ee wee 35. Rorippa. . F Plants pubescent with pews oo or stellate trichomes, and without unbranched o S. All “aaa uniformly ake. medifixed, appressed, sessile. 22. wi ioe himactay Seal new aca nls Me, Ol Shoe, dost ge ao ettech i ard es gos in ae obularia. S. Trichomes stellate, furcate, or peancneds) mixed with unbranched ones or not, stalked when stellate. Pr 1984] AL-SHEHBAZ, CRUCIFERAE 365 T. Petals deeply bilobed. nts scapose; fruits glabrous; ai of lateral stamens not appendaged; styles obsolete. ............ 24. Draba. U. Plants not scapose; fruits pene pubescent; filaments of lateral stamens appendaged; styles 1.5-4 mm long. ..... PRS Ree) 63, gst di Aa Sod dndem ohh eearanar deat MR 23. Berteroa. ale, eee entire or slightly emarginat V. Fru ae or oblong to rs often strongly com- ae Ww. ae wingless, biseriate, usually more than 4 per locule; fruits oblong or lanceolate. ............. 24. Draba. W. Seeds winged or margined, uniseriate, 1-4 per locule; fruits orbicular or nearly so X. Cauline leaves cuneate, not auriculate; styles less than | mm long; seed | per locule, strongly muci- laginous when wet. ............... 21. Alyssum. X. Cauline leaves auriculate; styles 1.5-3 mm as seeds 2—4 per locule, not mucilaginous when w BRS Gs). eiasn te eee OTs ews Ne an 25. aie V. Fruits ane or pyriform, inflated. Y. Fruits globose, not keeled, beakless; valve apex obtuse, aura at base of style; seeds usually orbicular; cotyle- dons accumbent. .................. 25. Lesquerella. Y. Fruits pyriform, keeled at the replum, ending abruptly in a stylelike beak; valve apex acuminate, extending 0.5— 1.5 mm into the beak area; seeds usually oblong; coty- ledons incumbent. .................. . Camelina. F. Fruits at least 4 times longer (often much more) than broad, linear or narrowly a. Plants with forked, branched, dendritic, or stellate trichomes, sometimes mixed with eee ones. b. Leaves 2- or 3-pinnatisect. ........ 0.0.0.0 ce eee 43, Descurainia. b. Leaves entire to ere lobed, never 2-pinnatisect. c. Stigmas with strongly decurrent lobes d. Stigma lobes with a lateral horn or + swelling in fruit. .......... eat Seach Ae eG eee Sh ek Ge Rr aera [Maithiola.] d. a lobes mauhout eu outgrowth. + g their entire length $s fruit; fruits sharply POUNLER Bi ty Fe A shiek eee [Malco mia. e. Biennials; stigma ieee free in mature fruits; fruits not a FR ed che x ere: a at Se oR PR er . Hesperis. Q . Stigmas capitate, san or 2-lobed (if 2-lobed, lobes shallow, diver- gent, never decurren f. Trichomes Seem medifixed, bifurcate mixed with 3-5-fur- cate or stellate ones, never unbranched; fruits mostly quadrangular LDESECTION yee eee ere ea grins seh Dace . Erysimum. f. Trichomes a mixture of 2 kinds: unbranched and stalked furcate; fruits terete or flattened parallel to the septum nt h. Plants not scapose; fruits mostly more than 2 cm long, more 366 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 than 10 times longer than broad; seeds sue a ar- ranged, very rarely obscurely biseriate. ...... Arabis. a. Plants glabrous or with unbranched trichomes only. i. Stamens long-exserted, equal in length; sepals strongly reflexed; petal claws papillose near the base; gynophores (3-)5-14 mm long. ....... 1. Stamens included or slightly protruding, tetradynamous or in 3 pairs of unequal length; sepals erect or spreading, very rarely reflexed; petal claws aa gynophores absent or very rarely present, to 3 mm long. its with a strongly developed beak in addition to style, rarely renal Saas (Diplotaxis and Erucastrum), cotyledons always conduplic k. Valves. a l pascal midnerve, with or without a few incon- spicuous later : (afi eae: ee fruits 4-angled. ...... 4. Erucastrum. l. Inflorescence ebracteate; fruits terete or flattened. m. Beaks strongly flattened, ensiform; stigma lobes decurrent: petals with dark brown or purple veins. ....... ruca. . Beaks conical or stylelike; stigmas entire or with nonde- current lobes; petal veins not darkly colored. n. Fruits flattened; seeds biseriately cae ovoid or ob- long, up to 1.5 mm long. ............. 7. Diplotaxis. n. Fruits terete; seeds uniseriately arranged, alobose, 1.5- n diameter. ................... 3. Brassica. k. Valves with 3-7 prominent nerves (these often more conspicuous n immature fruits). 0. Sepals erect, saccate at the base; petal claws as long as or longer _ = than the sepals. .......0.00.0.0 000.0000. eee, 5. Hutera. O. Nien widely Heer: or reflexed, not saccate; petal claws er than the sepals. .......0.0.0.0.0.0.0... 6. Sinapts. j. Fruits ene een accumbent or incumbent. Eras flattened parallel to septum; petals never yellow : om the base, coiling citcinately or spirally: replum flanged wits remains of valve seeds neither winged nor margined; plants perennials with ee or rhizomes, sometimes annuals or biennials. . Ua sui at eae e eAaiMunden sane Ae bedeed 27. Cardamine. q. Valves not dehiscing elastically, not coiling; replum margin no flanged; seeds winged or margined, very rarely wingless; are (ours) annuals or biennials. r. Cauline leaves all cera or pinnatifid. ...33. Sibara. r. Cauline leaves entire or toot s. Petals showy, purple to erm rarely lavender, crisped, 1-2 cm long, differentiated at base into an oblanceolate claw; sepals colored; buds acute; fruits on gynophores 1- DT OGG: ane etna sen Ye we teeta 2. Streptanthus. . Petals white or lavender, less than | cm long, not crisped, pit gradually attenuate to a clawlike i sepals green- ; buds obtuse; fruits sessile or nearly .32. Arabis. p. Fruits ae or quadrangular, very rarely cae totiened: petals commonly yellow, sometimes white or lavender t. Leaves entire, cordate-amplexicaul; mature fruits hae aie. seeds readily releasing abundant mucilage w Pai any bb Ave aed eee ep tana 4 ooh eed 13. ees nn 1984] AL-SHEHBAZ, CRUCIFERAE 367 t. Leaves (at least some of them) pinnately lobed or petiolate; fruits terete or obscurely 4-angled; seeds not mucilaginous (very rarely producing a thin coat of mucilage) when wet. u. Valves of fruits nerveless or obscurely nerved; seeds bise- riately or rarely uniseriately arranged; plants aquatic or 0 very wet habitats v. Flowers yellow; median nectaries present outside the in- ner stamens; leaves simple, sinuate or pinnately lobed; lower nodes usually lacking adventitious roots. ...... v. Flowers white or lavender; median neciatics eking: leaves pinnate; lower nodes with adventitious roots. beefy a doeenb easter tae ince oa eta 34. Nasturtium. u. Valves with a prominent midnerve; seeds uniseriately ar- ranged; plants mostly terrestrial. w. Lower leaves cordate or reniform, dentate, not auricu- late; seeds longitudinally striate. ........ 40. Alliaria. w. Lower leaves pinnate and/or auriculate; seeds reticulate. x. Flowers white or lavender; inflorescence an elongated raceme; cauline leaves serrate or entire, cuneate. 2A wad bitte, Seah tras lie Ate eget etna eet 29, Iodanthus. x. Flowers yellow; inflorescence corymbose; cauline leaves pinnately lobed, the uppermost sometimes un- lobed. y. Valves 3-nerved; fruits terete; cotyledons incum- bent; cauline leaves not auriculate............. . Sisymbrium. y. Valves |-nerved; fruits slightly flattened or some- what 4-angled; cotyledons accumbent; cauline leaves auriculate or amplexicaul. .. 28. Barbarea. —_— Full treatments of the tribes, including references, distributions, and aspects of their biology, will appear in separate papers as in the sequence below. The following brief accounts for the tribes are mainly intended to provide guidelines for the tribal limits and for the number of representative species and genera in the southeastern United States. Tribe 1. Thelypodieae Prantl in Engler & Prantl, Nat. Pflanzenfam. III. 2: 155. 1891 Herbaceous annuals or biennials, rarely perennials, glabrous or with simple hairs only; sepals equal at base or sometimes slightly saccate, erect or spreading to reflexed, occasionally forming an urceolate or bilabiate calyx; petals often strongly differentiated into claw and blade, usually crisped or channeled; sta- mens exserted or slightly protruding, equal in length or somewhat tetradyna- mous, rarely in 3 unequal pairs, not appendaged; siliques dehiscent, linear several to many times longer than broad, terete or flattened parallel to the septum, often borne on a distinct gynophore; seeds winged or wingless, not mucilaginous when wet; cotyledons accumbent or incumbent. (Including Stan- leyeae Robinson, Romanschulzieae O. E. Schulz, Streptantheae O. E. Schulz.) Type GeNus: Thelypodium Endl. 368 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 a L —VAVAAD seed, x 25; h, embryo, oriented as in seed, x 25; i, diagrammatic cross section of seed showing incumbent cotyledons, x 25. j, Coronopus didymus, fruit, x 6. k, Lepidium virginicum, fruit, x 6.1, L. campestre, fruit after removal of valve—note apical attach- ment of seed, x 6. m-t, BRASSICEAE. m, n, Brassica campestris: m, fruit, x 2; n, seed, x 6. o-q, Sinapis alba: o, fruit, x 2; p, embryo, x 6; q, diagrammatic cross section of seed showing conduplicate cotyledons, x 6. r, Diplotaxis muralis, fruit, x 3. s, Cakile edentula subsp. Harperi, fruit—note transverse joint, x 1. t, Calepina irregularis, fruit, x 6. petals removed, x 12; d, floral diagram; e, fruit, x 3; f, replum and septum, x 3; g, 1984] AL-SHEHBAZ, CRUCIFERAE 369 A natural tribe of 11 genera and some 110 species; represented in our area by seven indigenous species in two genera, Warea Nutt. and Streptanthus Nutt. Tribe 2. Brassiceae DC. Syst. Nat. 2: 152. 1821. Annual, biennial, or perennial herbs, sometimes subshrubs or large shrubs, glabrous or with simple hairs only; sepals erect or spreading, often saccate at base; petals usually clawed; stamens tetradynamous, the filaments very rarely appendaged; siliques long or short, dehiscent or indehiscent, often clearly dif- ferentiated into valvular and stylar (beak) segments, 1 or both seed-bearing, or the fruits transversely jointed or lomentaceous and breaking into parts, terete, angular, or flattened; seeds mucilaginous or not when wet, winged or wingless; cotyledons almost always conduplicate (very rarely accumbent or incumbent). (Including Cakilineae DC., Calepineae Godron, Erucarieae DC., Psychineae DC., Raphaneae DC., Velleae DC., Zilleae DC.) Type Genus: Brassica L. A natural tribe of some 52 genera and about 230 species; represented in the Southeast by 21 species, all (except four of Cakile Miller) naturalized weeds belonging to 11 genera: Brassica, Erucastrum Presl, Hutera Porta, Sinapis L., Diplotaxis DC., Eruca Miller, Raphanus L., Rapistrum Crantz, Cakile, Ca- lepina Adanson, and Conringia Heister ex Fabr. Tribe 3. Lepidieae DC. Syst. Nat. 2: 151. 1821. Annual, biennial, or perennial herbs, sometimes subshrubs or shrubs, gla- brous or usually pubescent with simple hairs only; sepals erect or spreading, rarely conspicuously saccate at base; petals often slightly differentiated into blade and claw; stamens 6, tetradynamous, or reduced to 4 or 2, the filaments frequently appendaged; siliques almost always shorter than 3 times their width, dehiscent or indehiscent, occasionally didymous and schizocarpic, always com- pressed at right angles to the septum (angustiseptate), replum much narrower than width of the fruit; seeds often mucilaginous when wet, winged or wingless; cotyledons accumbent or incumbent. (Including Brachycarpeae DC., Iberideae Godron, Isatideae DC., Senebiereae Godron, Subularieae DC., Thlaspideae DC.) Type Genus: Lepidium L A natural tribe of over 60 genera and more than 600 species; represented in our area by Lepidium, Coronopus Zinn, Cardaria Desv., Thlaspi L., Teesdalia R. Br., and Capsella Medicus, and about 16 species, of which all (except four species of Lepidium) are introduced weeds. Tribe 4. Alysseae Godron in Gren. & Godron, Fl. France 1: 112. 1848. Herbaceous annuals, biennials, or perennials, rarely subshrubs or shrubs, usually with branched or stellate trichomes, sometimes the trichomes simple or absent; sepals erect or spreading; petals attenuate at base or occasionally strongly clawed; stamens 6, tetradynamous, rarely 4, often with appendaged, toothed, or winged filaments, infrequently slender at base; siliques almost al- ways shorter than 3 times their width, dehiscent or rarely indehiscent, spherical, Ficure 2. Tribes Thelypodieae, Brassiceae, Pepidicne, 7 ee and Arabideae. a x — b, THELYPODIEAE, Warea Carteri: a, infructescence —note gynophor c-f, BRASSICEAE. c, Raphanus Raphanistrum, ee ee lomentaceous fruits, x 1h. d, R. sativus, fruit—note aborted lower segment, x 2. e, f, Eruca vesicaria subsp. sativa: e, fruit—note beak, x 2; f, fruit after fall of valves. g-l, Lepipiear. g-i, Thlaspi arvense. g, fruit, x 2; h, fruit after fall of valves, x 2; 1, seed, x 6. j-l, Teesdalia nudi- 1984] AL-SHEHBAZ, CRUCIFERAE | inflated, or commonly compressed parallel to the septum (latiseptate); seeds mucilaginous or not when wet, winged or wingless; cotyledons accumbent, ve rarely incumbent. (Including Camelineae DC., Drabeae O. E. Schulz, Lunarieae O. E. Schulz, Physarieae Robinson.) Type GENus: Alyssum L A tribe with poorly defined boundaries and comprising a heterogeneous assemblage of 41 genera and some 750 species; represented in our area by 20 species (13 indigenous) and seven genera: Lunaria L., Alyssum, Lobularia Desv., Berteroa DC., Draba L., Lesquerella S. Watson, and Camelina Crantz. Tribe 5. Arabideae DC. Syst. Nat. 2: 161. 1821. Annual, biennial, or perennial herbs, rarely subshrubs, glabrous or with simple, furcate, or branched hairs; sepals often ascending or spreading, equal or saccate at base; stamens tetradynamous, with slender or very rarely toothed filament bases; stigmas entire or slightly 2-lobed; siliques dehiscent, narrowly linear, sometimes oblong, rarely subspherical, often compressed parallel to the septum, occasionally inflated; seeds winged, margined, or sometimes wingless, cotyledons accumbent. (Including Cardamineae Calestani.) TyPE GENUus: Arabis L A tribe with a natural core of large genera and their relatives (but with artificially drawn boundaries) consisting of more than 570 species in 36 genera; represented in the Southeast by 44 species, of which only nine are introduced weeds, and ten genera: Cardamine L., Barbarea R. Br., Iodanthus Torrey & Gray ex Steudel, Leavenworthia Torrey, Selenia Nutt., Arabis, Sibara Greene, Nasturtium R. Br., Rorippa Scop., and Armoracia Gaertner, Meyer, & Scherb. Tribe 6. Hesperideae Prantl in Engler & Prantl, Nat. Pflanzenfam. III. 2: 154. Annual, biennial, or perennial herbs, rarely subshrubs or shrubs, glabrous or with simple, bifurcate, branched, or stellate trichomes, occasionally with multicellular glands; sepals erect, sometimes connivent; petals usually differ- entiated into blade and claw; stamens tetradynamous, filaments of the median pairs usually expanded at base or connate; stigma 2-lobed or very rarely entire, the lobes commonly decurrent; siliques long or short, dehiscent or rarely in- dehiscent, occasionally lomentaceous, infrequently beaked or appendaged; seeds often wingless; cotyledons incumbent or accumbent. (Including Anchonieae DC., Cheirantheae Villani, Erysimeae Reichenb., Matthioleae O. E. Schulz, Schizopetaleae R. Br. ex Barn.) Type GENus: Hesperis L. caulis: j, es x 6 k, fruit after fall of valves, x 6; 1, seed, x 12. m,n, ALysseAg, Lob- ularia maritit m, fruit, x 6; n, replum and septum, x 6. o-r, ARABIDEAE, Se/enia ured: 0, eee a x 1; p, seed—note wing, x 6; q, embryo, x 6; r, diagrammatic cross section of seed showing accumbent cotyledons, x 6. gz JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 i) cate AN Bee Ne L554} RE 3. Tribes Arabideae, Hesperideae, and Sisymbrieae. a—h, ARABIDEAE. a-<, ee a, L. sy Oe, plant with flowers—nole single-flowered scapes from center Liou bry tronalis, fruit, x 1; j, Er rysimum italy fruit, x 2. k-p, SIsyMBRIEAE. k—m, Alliaria petiolata: k, portion of plant w h flow x ‘a: 1, fruit, x 1; m, portion of fruit after removal of valve—note eee pean gh of seeds, x 3. n—p, Descurainia pinnata: n, tripinnatisect leaf, x '; 0, fruit, x 6; p, seed, x 25. 1984] AL-SHEHBAZ, CRUCIFERAE ae) A tribe with a fairly natural core of genera (but with highly artificial bound- aries) consisting of some 43 genera and about 500 species; represented in our area by six species in the genera Chorispora R. Br. ex DC., Hesperis, and Erysimum L. Tribe 7. Sisymbrieae DC. Syst. Nat. 2: 150. 1821. Annual, biennial, or perennial herbs, rarely shrubs, glabrous or with simple, furcate, or dendritic trichomes; sepals ascending or spreading; petals differ- entiated or not into claw and blade; stamens tetradynamous, the filaments linear and usually neither appendaged nor toothed; stigma entire or 2-lobed; siliques usually much longer than broad, dehiscent or rarely indehiscent; gen- erally terete or inflated; seeds primarily wingless, mucilaginous or not when wet; cotyledons incumbent. Type GENus: Sisymbrium L. A highly artificial tribe with some 70 genera and about 400 species; repre- sented in our area by seven species (one indigenous) in four genera: Alliaria Scop., Sisymbrium, Arabidopsis Heynh., and Descurainia Webb & Berth. ARNOLD ARBORETUM EN CAMBRIDGE, eee 02138 KAUL & ABBE, INFLORESCENCE ARCHITECTURE 375 INFLORESCENCE ARCHITECTURE AND EVOLUTION IN THE FAGACEAE Rosert B. KAUL AND ERNST C. ABBE INFLORESCENCE ARCHITECTURE is receiving increasing attention with respect to its role in the life of plants, but much remains to be learned of its effects on pollination and dispersal biology (see Wyatt, 1982). Little is known of the biology of fagaceous inflorescences or of their relationships with the growth or the reproductive patterns of the trees. Most studies (Abbe, 1974; Macdonald, 1979: Fey & Endress, 1983) have concentrated on the nature of the flower clusters (often called dichasia or partial inflorescences) and cupules. Hjelmqvist (1948) and Soepadmo (1972) briefly reviewed the variety of inflorescences in the Fagaceae. Celakovsky (1889) compared fagaceous inflorescences with bet- ulaceous and juglandaceous ones and enumerated the criteria by which he judged levels of specialization. Jager (1980) analyzed inflorescences of the Bet- ulaceae, the family perhaps closest to the Fagaceae. For this study, we have analyzed a sample of Fagaceae much larger than those of previous workers, and we here assess their inflorescences in terms of structural as well as functional criteria. Although our emphasis is on Paleotropical species, some species of the northern Temperate Zone are also included. The APPENDIX lists the species studied, as well as the provenance of the specimens. The family Fagaceae comprises perhaps 900 species. It is most richly rep- resented in eastern and southeastern Asia and the nearby islands of the western Pacific Ocean. In many places the family dominates the forests, and on a worldwide basis it is perhaps second in biomass only to the conifers (Soepadmo, 1972; Heywood, 1978). In tropical areas the Fagaceae are found from sea level to the frost line, but they are most abundant and often dominant at middle altitudes, where they favor rain forests of little seasonality. There the trees are evergreen and bear mostly entire to slightly sinuate or dentate leaves that sometimes have drip- tips. At higher latitudes in the Northern Hemisphere, the Fagaceae occur in both ever-wet and seasonally dry climates, where they are evergreen or decid- uous and their leaves are entire to sharply dentate or deeply lobed. The South- ern-Hemisphere genus Nothofagus grows in cool montane forests and is mostly evergreen. Castanea has about ten species in mesic forests of the eastern and southeastern United States, eastern Asia, southern Europe, northern Africa, and the Middle East. All are deciduous. The closely related Castanopsis has perhaps 120 species © President and Fellows of Harvard College, 1984. Journal of the Arnold Arboretum 65: 375-401. July, 1984. 376 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 in eastern and southeastern Asia and western-Pacific islands; there is a single species, C. chrysophylla, in montane forests of North America from Washing- ton to California. Lithocarpus (ca. 300 species) occurs from India to Japan, New Guinea, and Java. It, too, has a single North American representative, L. densiflora, in California and southwestern Oregon. Quercus (ca. 450 species) is widespread and often abundant across all the northern continents, as well as North Africa, at low and middle latitudes. It extends south in the Andean cordillera to about the equator. In Central Amer- ica, from the Isthmus of Tehuantepec to Colombia, there are 46 species (C. H. Muller, 1942), but in southeastern Asia, from Thailand to Java and Borneo, there are only 19 (Soepadmo, 1972). Trigonobalanus has three species: 7. verticillata Forman in southeastern Asia, southern China (Yunnan), and a few islands of the East Indies; 7. dai- changensis (A. Camus) Forman in Thailand; and 7. excelsa Lozano, Hernan- dez, & Henao in Colombia. Forman (1964) and Soepadmo (1972) recognized three subfamilies: Casta- neoideae (Castanea, Castanopsis, Lithocarpus s.1.), Quercoideae (Quercus s.1., Trigonobalanus), and Fagoideae (Fagus, Nothofagus). Crepet and Daghlian (1980) supported the close affinity of Quercus and Trigonobalanus, but Lozano and colleagues (1979) placed Trigonobalanus alone in Trigonobalanoideae. Kuprianova (1963) suggested that Nothofagus constitutes its own family, Noth- ofagaceae; Nixon (1982) concurred and also suggested betulacean affinities for it. Asiatic and western-Pacific Fagaceae have been variously interpreted at the species level by Camus (1929, 1934-1954), Barnett (1942, 1944), and Soe- padmo (1968, 1970, 1972). The most recent monograph (Soepadmo, 1972) is based on extensive field and herbarium work. Generic circumscription is complicated in Castanea, Castanopsis, Lithocar- pus, and Quercus by intermediate characters variously shared among them. Trigonobalanus is distinctive. Soepadmo (1972) has acknowledged the essential unity of Castanea and Castanopsis, the former differing from the latter only in having pistillate flowers always at the base of the spikes and bearing six or more stigmas. He rejected Hjelmqvist’s (1948) proposal of generic status (Chry- solepis) for the single species of western North American Castanopsis, but Forman (1966), Abbe (1974), and others have accepted it. Forman (1964), Soepadmo (1972), and Abbe (1974) recognized Quercus as distinct from Litho- carpus, despite historical confusion. They noted the constancy of differences including unisexual inflorescences, with the staminate ones pendent (Quercus), vs. bisexual erect inflorescences (Lithocarpus). There are floral differences as well. The distinctions between Lithocarpus and Castanopsis are usually obvious; except for a few species, the genera can be distinguished by the structure of their cupules (Forman, 1966; Soepadmo, 1972). The affinities of Fagus to the other genera—and even to Nothofagus—are less certain. In recent decades there has been no confusion concerning the identity of Fagus such as there has been about Castanea, Castanopsis, Litho- 1984] KAUL & ABBE, INFLORESCENCE ARCHITECTURE a7] carpus, and Quercus. These latter four genera form a coherent unit and, with Trigonobalanus, are the topic of this paper. Pollination in Fagaceae is by insects or wind. Castanea, Castanopsis, Litho- carpus, and Trigonobalanus verticillata are entomophilous, have scented flow- ers, and bear erect staminate inflorescences. Castanea and Castanopsis attract clouds of dipterans and other insects with their heavy, almost unpleasant odor; Lithocarpus is more delicately scented. Trigonobalanus daichangensis has pen- dulous staminate catkins and is anemophilous (Hjelmqvist, 1948; Soepadmo, 1972), as are Quercus, Fagus, and Nothofagus. Reports of entomophily in Quercus (e.g., Faegri & Wan der Pijl, 1979) are perhaps based on species of Lithocarpus placed in synonymy with Quercus. Trigonobalanus excelsa, re- cently discovered in Colombia (Lozano et al., 1979), is possibly entomophilous because it too has more or less erect staminate inflorescences. Except for the studies of entomophily in Castanea by Porsch (1950), Clapper (1954), and Jaynes (1974), there seem to be no definitive assessments of entomophily in the Fagaceae, however. The possibility that some entomophilous species are partially anemophilous (see Proctor & Yeo, 1972; Faegri & Van der Pijl, 1979) should be explored. Anemophily was present in the family by the Middle Eocene (Crepet, 1979) and was probably a factor leading to its prominence —especially that of Quercus — in north-temperate forests (Endress, 1977). Fossils from the Middle Eocene show floral structures almost identical to those of modern Castaneoideae (Cre- pet & Daghlian, 1980). Thus, the family was well advanced by the Middle Eocene and, in fact, is known from the Campanian and Santonian stages of the Cretaceous (J. Muller, 1981). The Fagaceae originated from unknown ancestors that were probably shared with Betulaceae and perhaps other families, but details are unclear. However, the Fagaceae form a large enough group—and one with enough diversity of inflorescence structure—to provide useful internal comparisons that suggest probable evolutionary pathways. MATERIALS AND METHODS We examined 126 species in Castanea, Castanopsis, Lithocarpus, Quercus, and Trigonobalanus, including thousands of specimens from our collections of Asiatic and western-Pacific Fagaceae as well as a few critical North American and European species. We and our colleagues collected in Paleotropical and eastern Asian forests, and for some species we have multiple collections from certain individual trees over a year’s time. Most of our Paleotropical specimens have been annotated by Soepadmo, and the nomenclature herein follows his treatment (Soepadmo, 1972). Each illustration represents an actual specimen at or near anthesis. Because the Fagaceae are so variable, no single figure can depict all the variance in a species or even on an individual tree. For illustration we have chosen species and individuals that best show the diversity of structure. Where possible, each figure shows current and previous flushes of growth. When leaves from the previous flush were present, as they often were (most JOURNAL OF THE ARNOLD ARBORETUM A é (ee ” head ci “, ce) 0. () eo 200000000- ° e @0000000. 0 ° oS 28 [VoL. 65 Ficures 25-28. Lithocarpus: 25, L. nieuwenhuisii; 26, L. caudatifolia; 27, L. curtisii: 28, L. kawakamii. and in that sense most of the tree is once part of an inflorescence. In some species the foliage leaves subtend individual spikes (e.g., Figures 11-13, 36, 40-45). The leader and most foliose branches are indeterminate. Indeterminate folio-bracteose branches are distally foliose and proximally bracteose (e.g., FiGures 29, 35, 39). Bracteose reproductive branches are often determinate (e.g., FIGURES 14, 16— 18). Those bearing staminate spikes and catkins are shed soon after anthesis, KAUL & ABBE, INFLORESCENCE ARCHITECTURE 385 ie} fo} ° ° ©} oe? ° ‘oO 0? ? ° i} * 0? 50n ° 6 8 %. b 0° 90° . 0 o fo) te ® 8 6 8 : "onrns O oO, e 3 is ° Py 0, 1 0 ‘0 ° fo 4 6 fa) o e Q, M806 ook ee 8 Aes “Po } °} ° 0-5 =o Q LJ bore } ° 9 Of or 0 ic bd ° fe) ° ° 3 ce) bre} %9 ° 2 Oo (6 9 ‘0, rs ° 3 0.99) 6 3 Og © ° o 9 0 0 } 20, ° g 0 367. ig 2 . 9, ° 9 06 6 oO Og 3 ¢ °o 96 3 re) ° 9° 9 0 ° 4 ° : ° 026 20, 7 5 Q c 7 5 005 ° 9 3 ° 3 A Co. o- 32 ° 000° ° 0 $ 9 o 5000S ° 2 oie) 2a 0 io 3 ° 9 q 06, G 3° Ogio 3) 0° 4 o §-0 90 > 90° 0° “ 28 9 8 8 90° 0° a) 09 re) fo) 0? ° °° re) 0° 0 0° ° oo is) 0° re} o? ° 60 a) xe) 0 40° fe) femme ° 0° 0° is} a0 ° o® ° ° 00 ° 200 ° ie} 200 ° Qo 0 5 2006 5 o 308 00 ee ) ° ° 9 OO000000000- VA VA 500020 9900 09° 00 -0090099% 400009 ° 9200006 fooe? 4000007 + 920009. o 9000° folohe) i) -9900°09 5 4090000 gor 900009 0° : : oe 00° 0° 29 02° 0° 0° 0° : o x) fe) Ko) 32 - 29, C. inermis; 30, C. fissa; 31, C. acuminatissima; 32, FiGures 29-32. Castanopsis C. rhamnifolia. while those with pollinated pistillate flowers are retained but usually drop their staminate spikes or flowers. These determinate branches are sometimes present on the same axis with indeterminate ones (e.g., FIGURES 22, 28, 31), in which case they are usually proximal and the indeterminate branches distal on the axis. Furthermore, they are often smaller than the distal branches and bear fewer spikes with fewer flowers. Sometimes sylleptic spikes and reproductive branches are accompanied by similar proleptic spikes and branches in a shoot system (e.g., FIGURES 16-19, 386 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 CASTANOPSIS 35 QUERCUS Ficures 33-39. Castanopsis and Quercus: 33, C. brevispina; 34, C. a 35, C. stipitata, 36, C. stellatospina; 37, 38, C. chrysophylla: 39, QO. uraia 30-32, 56). These proleptic structures are often strictly staminate, but the distal ones sometimes have a few pistillate flowers (e.g., FiGures 6, 16-19, 31, 41, 46 In some Fagaceae full repetition of the leader’s growth pattern, floral display, and sex distribution is obvious in the branches, whether they are sylleptic or proleptic (FiGures 10, 28, 31, 32, 41, 70A). In others the lateral branches are repetitive but determinate (FIGURE 26). More often, there is a basipetal gradient of decreasing similarity between the leader and its branches, both sylleptic and proleptic (FiGuRES 25, 27-31, 33-35, 70C, D). The most extreme manifestation of this gradient has small, determinate, bracteose branches bearing only sta- minate spikes or catkins (e.g., FiGuREs 25, 33-35, 44, 60, 70E-G). The middle branches, between the leader and the proximal branches, are frequently inter- 1984] KAUL & ABBE, INFLORESCENCE ARCHITECTURE 387 », CASTANEA Ficures 40-43. Castanea: 40, 41, C. mollissima; 42, C. crenata; 43, C. dentata. mediate in structure as well; they are often folio-bracteose, bear a few pistillate flowers, and may be determinate or indeterminate. The greatest differentiation between the leader and its branches occurs in Quercus, where in the most extreme cases some reproductive branches are distinctly short-shoots, are always proleptic and bracteose, bear only staminate catkins, and are entirely deciduous. The leader is foliose and bears pistillate spikes and sometimes also staminate catkins (FiGuREs 44, 47, 52, 66, 70F, G) Less extreme differentiation is shown by branches that have catkins axillary to the proximal leaves or bracts while the distal part of the axis is foliose, 388 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 QUERCUS ov O » 46 52 Ficures 44-52. Quercus: 44, 47, Q. glauca; 45, Q. championii; 46, Q. eas 48, Q. morii, 49, Q. phillyraeoides;, 50, QO. acuta; 51, Q. myrsinaefolia; 52, Q. gil\ sterile, and determinate or indeterminate (FiGuRES 46-49, 55, 56, 64, 65). Sometimes several degrees of differentiation between leader and branches exist on the same tree. DISTRIBUTION OF THE SEXES IN THE SHOOT SYSTEM Pistillate flowers, if present at all in a shoot system, are borne on distal spikes; the proximal spikes are staminate (FIGURES 6-13, 16-23, 25-52, 54, 56-58, 1984] KAUL & ABBE, INFLORESCENCE ARCHITECTURE 389 QUERCUS > snsarsvasesenredafeenssnnsennnens _ 62 6, QO. cambodiensis, 57, Q. mespilifolicides, 58, Q. griffithi ae = serrata, 60, Ficures 53-63. Quercus: 53, Q. kingiana; 54, Q. ae 55, Q. cf. chapensis; Q. championil, 61, Q. kinabaluensis: 62, QO. valdinervosa: 63, OG elt 62-66, 69, 70). Sometimes the leader has more pistillate flowers on more spikes than its branches (Ficures 7, 10, 30, 70A—C), but in other cases the distribution of pistillate flowers is about equal on leader and branches (e.g., Ficures 18, 26, 28, 31, 32). In any case, a clear acropetal gradient of increasing ““femaleness”’ is evident in almost every species examined. This gradient is variously ex- pressed in the leader, in the branches, and/or in the total inflorescence, and it complements the basipetal gradients of reduction of leaves to bracts and loss of indeterminacy in branches. In Quercus and Trigonobalanus daichangensis the sexes are almost always borne on separate axes (FIGURES 44-66, 69). Pistillate spikes (which sometimes 390 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 QUERCUS XN00, U0 4 a re) O fe} ie} SN erst sf & ° ° PS 9 OF fe) 3 ° a ° ie) re} fe} / Oo 3 ° O° fo - 3° oO 3 ° ce) 3)8 Les 82/88 a ase: ° ° 8/8 $5608 SB 9s s 88938 a 0699 ° fe) rome) 03 9 9000 9 9 OOF a2 He 2000 ° 052 ° 090) _ 4 0208 520 ° ot 028 O20 ° fe) O88 oo oO MO OO en more) 8oF 58 98 328 8 0° 9 oO oo (e) oo 6 64 Poo? 5 66 TRIGONOBALANUS a e e e e Pid Lo e Q S . ° e ca 7 ° e “4 oC fo} ie) fe] % Le} 00 . wey: fore) ee fe} c Cc iS c Cc oO ee ie) fe} oO Oo Oo ° ° ce} fe} ° 68 691° 67 Ficures 64-69. Quercus and Trigonobalanus: 64, Q. imbricaria, 65, 66, Q. borealis; 67, 68, 7. verticillata, 69, T. daichangensis (after Forman, 1964). exist as a single cymule or flower) are invariably distal and are nearly always subtended by a leaf. Staminate catkins occur singly in the axil ofa leaf or bract, mostly in the proximal part of the shoot system (Figures 44-53, 55-59, 61, 64-66, 69). They are shed soon after anthesis. Catkin-bearing short-shoots are proleptic (e.g., Figures 44, 47, 48, 52) and are also shed soon after anthesis. In Trigonobalanus verticillata some shoots bear only staminate spikes, while others have androgynous ones (FiGuRES 67, 68); our specimens and observa- tions are inconclusive as to the existence of a sex gradient in the total inflo- 1984] KAUL & ABBE, INFLORESCENCE ARCHITECTURE 39] rescence. Likewise, information about sex distribution in 7. exce/sa is ambig- uous. According to Lozano and colleagues (1979), staminate and pistillate inflorescences are borne separately on the tree. In the northern genus Castanea (FiGures 40-43) the ratio of staminate to pistillate cymules is higher than in Castanopsis or Lithocarpus. However, the only two American species of the latter two genera, Castanopsis chrysophylla (Ficures 37, 38) and Lithocarpus densiflora (FIGURE 15) also have such a higher ratio, and they too are northern. This ratio apparently also exists in Quercus, where the northern species bear fewer pistillate cymules on a branch than do many tropical species. SYLLEPSIS OF THE SPIKES The spikes contain two kinds of axes: 1) an elongate primary axis or long- shoot to which the 2) cymose short-shoots are racemosely attached. Second- order sylleptic long-shoots are rather common in the staminate spikes of Litho- carpus and Castanopsis chrysophylla (FiGures 7, 21-24, 37, 38, 67) and are occasional in Castanea. An occasional bisexual spike also has sylleptic branches (FIGURE 23). Such sylleptic branching produces ramified spikes with branches at what would otherwise be sites of cymules and each second-order long-shoot is subtended by a primary bract in the proper phyllotactic position (FIGURE 2). The truly sylleptic nature of this branching 1s evidenced in FiGurE 2, where second-order branches are appearing on an immature primary axis. Sylleptically branched spikes differ from bracteose reproductive branches (as defined above) in that they bear dichasia on the primary axis and terminate in a floriferous axis. There is no functional or abortive bract-clad apex of the kind found on typical vegetative branches. In some instances, typical bracteose reproductive branches bear a distal spike that overtops the primary axis (see FiGuRE 26), and the abortive apex is then apparently lateral. Although such a branch appears to bea sylleptically branched spike, it is readily distinguished from a truly sylleptic spike by the large, lateral, bract-clad abortive apex. DISCUSSION Studies of floral development and morphology have produced a large body of literature on evolutionary pathways and adaptations in flowers. Although literature concerning inflorescences is less definitive, many of the same forces behind floral evolution are clearly also involved in inflorescence evolution, with analogous morphological results. Flowers and inflorescences share such major functions as effecting pollination and nurturing and disseminating seeds and fruits. Such well-known floral evolutionary phenomena as pleiomery, oligomery, heterochrony, changes in size and symmetry, protogyny and protandry, and transfer of function are also found in inflorescences. We can therefore apply some of the principles of floral evolution to inflorescen In the history of plants, vegetative and reproductive fonetaus became sep- 392 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 arated and strobili, flowers, and inflorescences were produced. Further phy- logenetic differentiation in flowers was by continued separation of functions, producing such advanced features as imperfect flowers, dioecy, anemophily, or elaborate forms of entomophily, and a host of attendant morphological features. These have appeared repeatedly and independently in diverse angio- sperms responding to similar evolutionary pressures. Apparent reversal of these trends, such as anemophily reverting to entomophily, involves further modi- fications of existing structures and behaviors in response to new evolutionary pressures similar to ancestral ones. There is saan and ecological evidence that the outcrossing produced by many of these structures and pollination systems is advantageous to the species. Temporal ree spatial separation of the sexes is most evident in highly evolved plants such as the Fagaceae. In the Fagaceae some evolutionary patterns have recurred at successively higher levels of morphological organization. The separation of sexes into dif- ferent flowers, which is almost total in extant Fagaceae, preceded separation of imperfect flowers on the spike, of staminate from pistillate spikes on the same shoot, and of shoots bearing only staminate or pistillate spikes. Many stages of these events occur in various combinations in living Fagaceae, but of the genera studied, only Quercus and a few species in other genera show them in their fullest expression. Paralleling these events are the beginnings of protandry at the inflorescence level. This is a natural consequence of the acropetal ontogeny of shoots and spikes, the acropetal anthesis of flowers, and the restriction of female flowers to distal regions of the shoots. None of the genera studied, however, is mor- phologically dioecious, although some may approach functional dioecism. The frequent lack of synchrony among adjacent conspecific individuals (which we often saw in the field) contributes another dimension to incipient functional dioecism. The Fagaceae also show stages of separation of reproductive from vegetative functions. Foliose reproductive branches are usually indeterminate and perform all functions, including extension growth. Merely bracteose reproductive branches are mostly determinate and abscise quickly if they bear only staminate spikes, but they are determinate to indeterminate if they have pistillate flowers. Compared with tropical species of Fagaceae, more northerly members show reduced or, most often, no syllepsis in their shoots or spikes. (The cymulose short-shoots are, of course, sylleptic on their long-shoots, as are spikes on the shoots that bear them.) The spikes of Castanopsis as ysophylla and mee densiflora (and occasionally Castanea) show it only weakly. Even within tanopsis and Lithocarpus, there is probably a general loss of syllepsis as one moves from the tropics into the Northern Hemisphere, while the strictly ex- tratropical Castanea has nearly lost it altogether. In Quercus there is none in the catkins of any species we studied, and only one of our species, Q. kingiana from Burma (FiGuRE 53), showed vegetative syllepsis. These observations are consistent with the well-known decline of syllepsis in trees and shrubs in general as one moves northward from the tropics. It 1s thus possible that syllepsis is an ancestral condition in the Fagaceae and other 1984] KAUL & ABBE, INFLORESCENCE ARCHITECTURE os % 0, 0. cae *eN “ON DUNS > me) a *9000000000000: o co) oo ce) ° ° 6 ° oO ‘o o 5 io) on > o, oO 2000000000000) ° © 3 ° > s. 5 Ko > . © oy 'e 0. cs) ss 9900000000 ° % fey . ‘0 Q% % o, i )A0O00000000: 0 ” 393 Q a or 00 oe re) °° oI Aw, 00000000000. 4s o 0000000000: ‘. $00000000000. ~ 990000000000 Figure 70. Possible derivations of inflorescence patterns in Lithocarpus, Castanopsis, Castanea, and Quercus (downward-pointing arrows indicate deciduous branches). 394 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 angiosperms, and that it was often reduced or lost in woody plants as they colonized cooler climates. Figure 70 summarizes logical levels of specialization, based upon the con- cepts advanced above, in modern species of Castanea, Castanopsis, Lithocar- pus, and Quercus. No hypothetical or extrafamilial patterns are shown. The coexistence of ancestral and derived characters among living members of such a large family is to be expected, although even the least derived stages (FIGURE 70A, B) are quite specialized for angiosperms. All the stages shown, as well as numerous intermediates and variants, are common in living Fagaceae and must be considered evolutionarily successful. FiGuRE 70A, B shows flowers of both sexes on some or all branches as well as on the leader—distributions common in Lithocarpus and Castanopsis. Some perfect flowers occur but are not indicated. Both illustrations also show pro- leptic and sylleptic indeterminate branches. The sylleptically branched sta- minate spikes of Figure 70B have perhaps preceded the condition in FIGURE 70C, where sylleptic branching is lost in both the leader and the spikes, and fertile branches are indeterminate with at least some exclusively staminate. Spikes are seldom ramified, and pistillate flowers are distal on the shoots but proximal on the spikes. This pattern occurs in Castanea (Ficures 40-43) and some species of Castanopsis. FiGure 70D is similar to 70C, but the fertile portions of the branches are bracteose rather than foliose. Extension growth produces normal foliage leaves. The spikes are all unisexual. This pattern occurs in Lithocarpus (FIGURE 28) and Castanopsis (FIGURE 35). In Figure 7OE the branches are all proleptic and staminate. The upper branches are indeterminate, and the fertile region is bracteose. Lower branches are determinate and usually caducous. This combination of fertile determinate and indeterminate branches occurs in Castanopsis and Lithocarpus. A common condition in Paleotropical Quercus is shown 1n FIGURE 70F. The leader has mostly pistillate spikes, and it sometimes has a few staminate catkins as well. Indeterminate branches can bear pistillate spikes and often have sta- minate catkins toward the base. Short-shoots bearing catkins are borne in leaf axils and are soon abscised. The most extreme condition in extant Fagaceae is shown in FiGuURE 70G and is commonest in northern Quercus. The pistillate spikes have one or few flowers. The leader lacks staminate catkins or has only a very few. The proximal branches are determinate, caducous short-shoots, and the distal branches are indeterminate. There are many variants of this and the preceding pattern. The verticillate phyllotaxy and resultant verticillate display of inflorescences in Trigonobalanus verticillata is unique in the family, although some Quercus species produce crowded and essentially verticillate leaves near the end of a flush. Furthermore, 7. verticillata produces alternate and then opposite leaves on many branches before forming whorled leaves with axillary inflorescences. Forman (1964) believed the pendent catkins of Quercus and Trigonobalanus daichangensis to be derived from erect spikes of entomophilous Fagaceae, an opinion shared here and consistent with that of Schottky (1912). Forman (1964) also interpreted the single flowers on spikes and catkins of some Quercus and 1984] KAUL & ABBE, INFLORESCENCE ARCHITECTURE TABLE 1. Probable evolutionary trends in fagaceous inflorescences. CHARACTER ANCESTRAL CONDITION DERIVED CONDITION — . Vegetative branching sylleptic and proleptic only roleptic 2. Branches indeterminate, determinate, foliose bracteose 3. Spike- and catkin-bearing branches persistent caducous 4. Flowers of spike-bearing branches staminate and staminate or pistillate istillate 5. Flowers of leader staminate and pistillate only pistillate 6. Spikes sylleptically unbranched branched 7. Flowers of spikes staminate and staminate or pistillate or pistillate perfect 8. Staminate flowers eas in spikes catkins 9. Pistillate flowers on s numerous few or one 10. Pistillate flowers in ai inflorescence numerous few to none 395 Trigonobalanus species as having been derived from dichasial clusters. 77i- gonobalanus verticillata and T. excelsa have dichasial flower clusters, while 7. daichangensis has both clusters and single flowers. In addition, 7. daichangensis has pendent catkins and is anemophilous, but the other two species have more or less erect male spikes and are probably entomophilous. Thus, 77rigonobal- anus is transitional between the entomophily of the Castaneoideae and the anemophily of Quercus. Evolutionary increase in flower numbers is likely in some cases, especially in Lithocarpus, Castanea, and Castanopsis. On the other hand, there are fewer female flowers per spike and per total inflorescence in many northern Fagaceae. Catkins and spikes of Quercus generally bear fewer flowers than comparable axes of the other genera studied. Furthermore, there are reductions in floral and dichasial morphology in Quercus. The catkin can be interpreted as the least expensive structure, in terms of pee and materials, that can be produced quickly at the start of the relatively sh growing season. In addition, Quercus species 1n northern ee are conspecifically more i ame and usually have fewer congeners than do species of Lithocarpus stanopsis, and Quercus in the tropics. In the north Quercus shows strong mes conspecific pollination synchrony, while tropical Lithocarpus and Castanopsis (and perhaps Quercus, too) conspecifics frequently show marked reproductive asynchrony. The likelihood of successful pollination is thus perhaps as great in northern Quercus as in the other genera, which have more numerous flowers and are usually scattered in species-richer populations. The presence of relatively few species of Quercus in the lowland Paleotropics suggests that members of the genus have not been especially successful in migrating to warmer from cooler regions, where they probably originated. The converse is probably the case 396 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 with Lithocarpus and Castanopsis, both of which have fewer temperate than tropical species. TABLE | summarizes likely evolutionary trends in fagaceous inflorescences, as portrayed in FiGure 70. Intermediates and combinations of these characters can be found in many species and sometimes on a single individual. Within flower and fruit characters, ecology, and geography, there are correlates to these inflorescence characters but our data are as yet inconclusive. We suspect that inflorescence characters and fruit size are related. The range in dry-fruit weight in our Paleotropical specimens is from less than 0.5 to more than 60 g. A few extant Fagaceae bear pedunculate pistillate cymules (e.g., FiGures 1, 6, 8) that are perhaps indicative of ancestral thyrselike conditions. Soepadmo (1972) considered Lithocarpus elegans (Blume) Hatus. ex Soep. to be probably the most primitive in its genus. It has staminate cymules with up to 24 flowers, and pistillate ones with up to 10; the staminate spikes are frequently much branched. Hjelmqvist (1948) regarded Lithocarpus as the primitive genus in the family based on characters of the flowers and cymules. Our observations of inflorescences support that conclusion. The Betulaceae are sometimes cited as the closest living relatives of the Fagaceae. Abbe (1935) showed the possible derivation of betulaceous inflo- rescences from thyrselike predecessors, a pattern perhaps similar to that of the Fagaceae. Jager (1980) analyzed betulaceous inflorescences and postulated re- duction in number and size of catkins, increase in winter protection of catkins, and reduction of leaf size, all associated with adaptation to colder climates with shorter growing seasons. The more primitive Betulaceae occur in warmer Sino-Himalayan areas. These attributes parallel those of the Fagaceae to some extent, but the Betulaceae are generally more northern and lack large numbers of tropical or subtropical species that might be of value for comparison. Further, there are no entomophilous Betulaceae. Many inflorescence characters of the Fagaceae parallel those found by Man- ning (1938) in the Juglandaceae. That family exhibits trends 4-10 shown in TABLE |. There are also geographic correlations of structure that point strongly to congruent biogeographic histories of the two families. Celakovsky (1889) analyzed inflorescence patterns in the Fagaceae, Betula- ceae, and Juglandaceae and found strong tendencies for proleptic shoots to be reproductive and bear reduced leaves. He interpreted single perfect flowers in leaf axils as primitive in these families’ ancestors. In our study we have applied the principles of differentiation used by Celakovsky¥, with some modifications. We believe that he was correct in his assessment of fagaceous inflorescences, although his conclusions were based on only a few species. Further, we agree that the single perfect axillary flower is the primitive condition for angiosperms. The flower as a reproductive short-shoot is but one of a series of evolutionary repetitions of the tendencies to concentrate and elaborate reproductive struc- tures ACKNOWLEDGMENTS This study was funded by National Science Foundation grants DEB-7921641 and DEB-8206937 to R. B. K. Fieldwork was supported by N. S. F. grants 1984] KAUL & ABBE, INFLORESCENCE ARCHITECTURE sof G-6369 and G-15926 to E. C. A. We are especially grateful for the field and laboratory help of Lucy Boothroyd Abbe, and for the assistance of J. A. R. Anderson, A. L. Bogle, M. P. Bogle, H. M. Burkill, E. Ensor, H. B. Gilliland, M. L. Kaul, H. Keng, T. Koyama, W. Meijer, L. Metcalfe, G. Mikal, J. V. Pancho, B. Rollet, J. S. Sinclair, T. Smitinand, B. E. Smythies, E. Soepadmo, H. C. Tang, P. Wardle, J. S. Womersley, and J. Wyatt-Smith in collecting, identifying, and preparing specimens. We are indebted to the staffs of a, BH, BKF, BM, BO, DD, E, GH, K, KEP, L, LAE, MIN, PNH, SAN, SAR, SING, TI, and us for their assistance during our visits. Helpful comments on the manuscript were provided by M. R. Bolick. LITERATURE CITED Aspe, E.C. 1935. Studies in the phylogeny of the Betulaceae. I. oe and inflorescence anatomy and morphology. Bot. Gaz. (Crawfordsville) 97: 1-67. 1974. Flowers and inflorescences of the * ae ” Bot. Rev. 40: 159- B61 Barnett, E. C. 1942. The Fagaceae of Thailand and their geographical distribution. Trans. Bot. Soc. Edinburgh 33: 327-343. 1944. Keys to the species groups of Quercus, Lithocarpus, and Castanopsis of eastern Asia, with notes on their distribution. /bid. 34: 159-204 pens B.G., & L. A. S. JoHNson. 1979. Evolution in the ee from inflorescence structure. Proc. Linn. Soc. New South Wales 102: 157- Camus, A. 1929. Les chataigniers. Monographie des genres Castanea - : ‘aslanousis: sae ee Sylvic. 3: 1-604. 4-1954,. Les chénes. Monographie du genre Quercus. Ibid. 6: |-686; 7: 1- 830: Fi 1-1314 CELAKOVSKY, L. 1889. O fylogenetickém vivoji rostlin jehnédokvétych. Sitzungsber. KGnigl. B6hm. Ges. Wiss. Prag, Math.-Naturwiss. Cl. 1889: 319-343. CLapper, R. 1954. Chestnut breeding, Peat and results I. Breeding material and pollination techniques. J. Heredity 45: 106-11 Crepet, W. L. 1979. Some aspects of the ee biology of Middle Eocene angio- sperms. Rev. Paleobot. Palynol. 27: 213-238. & C. P. DAGHLIAN. 1980. Castaneoid inflorescences from the Middle Eocene of Tennessee and the diagnostic value of pollen (at the subfamily level) in the Fagaceae. Amer. J. Bot. 67: 739-757 Enpress, P. K. 1977. ies trends in the Hamamelidales-Fagales-group. PI. Syst. Evol. Suppl. 1: 321-34 Faccri, K., & L. VAN DER au ‘1979, The principles of pollination ecology. ed. 3. Pergamon, New York Fey, B. S., & P. K. ENpress. 1983. Development and morphological interpretation of the ety in Fagaceae. Flora 173: 451-468. ForMAN, L. 1964. aati a new genus of Fagaceae, with notes on the ee of the family. Kew Bull. 17: 381-396. 19 Generic ea. in the Castaneoideae (Fagaceae). /bid. 18: 421- fidie. A R. A. A. OLDEMAN, & P. B. TOMLINSON. 1978. Tropical trees and forests. Springer-Verlag, New York. Heywoop, V. H., ed. 1978. Flowering plants of the world. Mayflower, New York. Hyetmovist, H. 1948. Studies on the floral morphology and phylogeny of the Amen- tiferae. Bot. Not. Suppl. 2: 1-171. JAGeR, E. J. 1980. Progressionen im Synfloreszenzbau und in der Verbreitung bei den Betulaceae. Flora 170: 91-113. 398 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 65 Jaynes, R. A. 1974. Genetics of chestnut. U.S. Forest Serv. Res. Pap. WO-17: i-1v, -13. a L. A. 1963. Ona hitherto undescribed family belonging to the Amen- e. Taxon 12: 12, 13. es C. G., J. HERNANDEZ-CAMACHO, & J. E. HENAo-S. 1979. Hallazgo del género Trigonobalanus Forman, 1962 (Fagaceae) en el Neotr6pico—I. Caldasia 12: 517- 537. MacpbonaLp, A. D. 1979. Inception of . cupule of Quercus macrocarpa and Fagus i ae Canad. J. Bot. 57: 1777-1782. MANNING, W 1938. The eee of the flowers of the Juglandaceae. I. The ee Amer. J. Bot. 25: 407-419. Mutter, C. H. 1942. The Central American species of Quercus. U.S.D.A. Misc. Publ. 477. U.S. Govt. Printing Office, Washington, D. Mutter, J. 1981. Fossil pollen records of extant angiosperms. Bot. Rev. 47: 1-142. Nixon, K.C. 1982. In support of recognition of the family Nothofagaceae Kuprianova. Bot. Soc. Amer. Misc. Publ. 169: 102 Porscu, O. 1950. Geschichtliche Lebenswertung der Kastanienbliite. Oesterr. Bot. Z. : 269-321. Proctor, M., & P. Yeo. 1972. The pollination of flowers. Taplinger, New York. SCHOTTKY, E. 1912. Die Eichen des eer ae gee Ostasiens und ihre pflanzengeo- aphische Bedeutung. Bot. Jahrb. Syst. 47: 617-708. een E. 1968. A revision of the genus Quercus L. subgen. Cyclobalanopsis (Oer- = Schneider in Malesia. Gard. Bull. Singapore 22: 355-427 70. Florae Malesianae Seg alia XLIX. Malesian species of Lithocarpus BL. oo Reinwardtia 8: 8. . 1972. Fagaceae. FI. ae : 712): 265-403. Troi, W. 1964. Die Infloreszenzen. Gustav Fischer, Stuttgart. Wyatt, R. 1982. Inflorescence architecture: how flower number, arrangement, and phenology affect pollination and fruit-set. Amer. J. Bot. 69: 585-594. APPENDIX. Species studied and provenance of specimens. CASTANEA C. crenata Sieb. & Zucc. C. dentata Bor C. mollissima Alamé CASTANOPSIS Taiwan (cultivated) Nebraska (cultivated) Nebraska (cultivated) javanica (Blume) A. D - lanciifolia (Roxb.) Hickel & A. Camus inermis (Lindley) Bentham & Hooker f. C. acuminatissima (Blume) A. DC. Vietnam C. argyrophylla King Thailand C. armata Spach Thailand C. brevispina Hayata Taiwan C. chevalieri Hickel & A. Camus Vietnam C. chrysophylla (Douglas) A. DC. California, Oregon C. costata (Blume) A. DC. Borneo (Sabah) C. cuspidata Schottky Japan C. le eel A. DC. Thailand C. ferox Spa Thailand C. fissa ( ae Rehder en Wilson Hong Kong C. formosana (Skan) Hay aiwan C. foxworthyi Schottk Borneo (Sabah) C. C. C aya Indonesia (Java) Thailand 1984] C. C.m C. C. C. C. Cc. C. pu C. longipetiolata Hickel & A. Camus egacarpa Gamble motleyana King nephelioides King ex Hooker f. philippinensis (Blanco) Vidal cf. pierrei Hance pyriformis nis & A. Camus urpurea Barn rhamunifolia aan A. DC. C. schefferiana Hance c, C stellatospina Hayata stipitata Hayata C. subacuminata Hayata Cc ae tribuloides (Smith) A. DC. tungurrut (Blume) A. DC. LITHOCARPUS Beene ee a ee ee NUE Ee ele . aggregata Barn . amygaalifolia (Skan) Hayata bennettii (Miq.) a er . brevicaudata ia . buddii (Merr.) A. Camus . cantleyana (King) Rehder ’ caudatifolia (Merr.) Rehder . celebica (Miq.) Rehder . clementiana (King ex Hooker f.) A. Camus . curtisii (King ex Hooker = A. Camus . dasystachya (Miq.) Rehd . dealbata (Hooker f. & TH oRisen) Rehder . densiflora (Hooker & Arn.) Rehder ee A. Camus . edulis Naka . elegans (Blume) Hatus. ex Soep. . ewyckii (Korth.) Rehder . fenestrata (Roxb.) Rehder . formosana Hayata . garrettiana (Craib) A. Camus . gracilis (Korth.) Soep. . hancei (Bentham) Rehder . harmandii (Hickel & . eee A. Camus . havilandii (Stapf) Barn . hendersoniana A. an . hystrix (Korth.) Rehder . kawakamiui Hayata . kodaihoensis Hayata . konishii (Hayata) Hayata . lampadaria (Gamble) A. Camus . leptogyne (Korth.) Soep. . lucida le ) Rehder macphailii (M. R. sey ee Barnett . maingayl an Rehde KAUL & ABBE, INFLORESCENCE ARCHITECTURE Cambodia alaya Borneo (Sarawak) Malaya Philippines hailand Vietnam Thailand Thailand Burma Indonesia (Java) Thailand Taiwan Borneo (Sabah) Cambodia Taiwan eset Sing eee (Sabah), Philippines Philippines Borneo (Sarawak) Hong Kong Borneo (Sabah, Sarawak) Burma, Thailan California, Oregon Vietnam Japan hailand Borneo (Sabah) rma Malaya 399 Q JOURNAL OF THE ARNOLD ARBORETUM meljeri Soep ee (Elmer) Rehder nantoensis Koidz. neorobinsonit A. Camus nieuwenhuisti (Seemen) A. Camus _ papillifer Hatus. ex Soep. . pattaniensis Barnett pees (Wallich) Rehder assa (Miq.) Rehder ed A. Camus rhombocarpa Hayata rufovillosa Soep. . soleriana (Vidal) Rehder Sues (Craib) A. Camus SUN a (Blume) Rehder . ter a upula Hayata . thomsonii (Miq.) Rehder . truncata (King ex See f.) Rehder & Wilson . turbinata (Stapf) F urceolaris (Jack) Mer. wallichiana (Lindley ex Hance) Rehder woodii (Hance) A. Camus wrayi (King) A. Camus ERCUS . acuta Thunb. . borealis Michaux . brandisiana Kurz . cambodiensis Hickel & A. Camus un - griffithii Hooker f. & Thomson imbricaria Michaux Lecaieoa Soep. sill . lana 2 Smith one Kin merrillii Seemen mespiifolicides A. Camus mort! Hayata . myrsinaefolia Blume . pachyloma Seemen . paucidentata Franchet . phillyraeoides A. Gra . salicina Blume serrata _ stenophylloides Hayata . Subsericea A. Camus uraiana . valdinervosa Soep. . virginiana Miller x se) << > Borneo (Sabah) Philippines Malaya Borneo (Sabah) Borneo (Sabah) Borneo (Sabah) Malaya Thailand New Guinea Philippines Thailan Borneo (Sarawak) Taiwan Thailand Burma Borneo (Sabah) Borneo (Sabah) Philippines Thailand Japan Nebraska Burma, Thailand Cambodia Hong Kong Oregon Borneo (Sabah) Japan, Taiwan Taiwan Thailand I Thailand ap ae ene Tai Borneo (Sabah) Borneo (Sabah) Florida [voL. 65 1984] KAUL & ABBE, INFLORESCENCE ARCHITECTURE 401 TRIGONOBALANUS T. verticillata Forman Borneo (Sabah) R. B. K. E.C.A. SCHOOL OF BIOLOGICAL SCIENCES DEPARTMENT OF BOTANY UNIVERSITY OF NEBRASKA UNIVERSITY OF MINNESOTA LINCOLN, NEBRASKA 68588-0118 St. PAUL, MINNEsoTa 55108 WHITE, ARALIA SPINOSA 403 THE ARCHITECTURE OF DEVIL’S WALKING STICK, ARALIA SPINOSA (ARALIACEAE) PETER S. WHITE ARALIA SPINOSA L., a small tree of moist sites in eastern deciduous forests, is striking in appearance. Among its distinctive traits are large, two- or three- times compound leaves (often over | m long and with more than 100 leaflets), abundant prickles, large terminal inflorescences (over 6000 flowers), and a relatively unbranched pachycaulous habit. Several of these characteristics are treated in the empirical observations known as Comer’s rules (Hallé et al., 1978), which state that the thicker the annual increment, the larger the leaf borne on the increment and the fewer the ultimate number of branches. A test of Corner’s rules in eastern deciduous trees (P. S. White, 1983) confirmed that Aralia is extreme in form: it had larger leaves, thicker annual increments, and fewer branches per unit leaf area than any of the other 47 species considered. The ecological role of Aralia is also of interest. Although trees of this species sometimes persist in shade, they occur primarily in disturbed areas (where they appear after fire, windstorm, or gap creation) and in old fields. The relationship between the architecture and the ecological role of a tree has been the subject of a growing literature (e.g., Horn, 1971; Whitney, 1976; Givnish, 1978a; Hallé et a/., 1978). Aralia was selected for study bec: several ofits morphological traits (large lincrement (Marks, 1975), high numbers of leaves and extensive total leaf area per annual increment (P. S. White, 1983) and larger leaf size with compound organization (Givnish, 1978a, 1978b)) have been hypothesized to be correlates of shade intolerance. In this study I investigated the gross morphology and life history of Aralia ramets. Of primary interest were the growth in height and in diameter of individual ramets, the accumulation of branches in Aralia crowns over time, and the leaf area carried by these branches. METHODS Three populations of Aralia spinosa occupying moist, low-elevation (500- 600 m) sites in the Great Smoky Mountains, Sevier County, Tennessee, were sampled. These populations were irregular in extent due to past agricultural disturbances. Since Aralia spinosa is clonal, individual stems are ramets; genets were not distinguished here because this would have greatly disturbed the populations. A random-numbers table was used to sample 114 ramets—one- fourth of all stems encountered. Because leaf scars, terminal bud-scale scars, and inflorescence axes all persist © President and Fellows of Harvard College, | Journal of the Arnold Arboretum 65: 403-418. duly; 1984. 404 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 for ten to fifteen years, the developmental history of each ramet could be reconstructed. For each year of growth, the amount of extension growth (the annual increment, or the length of axis produced per year), the number of were recorded. Analysis of architectural development was facilitated because flowering and branching are intimately related: each flowering event terminates the growth of the axis, with branching occurring the following year from axillary buds that develop below the old inflorescence axis. Each year’s increment was numbered starting at the base of the ramet (annual increment no. |) and working to the tips of the branches (the most recent year’s growth) (see Figure 1). Yearly increments were also numbered from the base of each branch order. In this way, the increment formed each year could be analyzed as a function of age and crown position. The numbering system was used to divide annual increments into trunk increments (“T” in the figures; the unbranched parts of the ramet) and first- and subsequent-order branch increments (“B” in the figures). Spur branches (“Sp”; short, slow-growing, suppressed branches) were so morphologically dis- tinctive that they were segregated from other branches in the analysis, as were inflorescence axes (““F” Part of one population had been cut during roadside mowing the previous season (1980). The trunks that sprouted after this grew more vigorously than did uninjured trunks. Shaded stems, on the other hand, grew less vigorously than did uninjured ones. Hence, for the trunk-building phase of growth, three states could be contrasted: injured, open-grown (“I’’ in the figures), uninjured, open-grown (“T’’), and uninjured, shaded (‘‘S’’). For comparisons between trunks and branches, only open-grown, uninjured trunks (““T’’) were used. Note that the “T” Noe Lae unsuppressed trunks) in the figures was used twice: once in the “T” vs. ““B”’ (branches), ““F’’ (inflorescences), and “‘Sp”’ (spur branch- es) comparison, ina again in the “T”’ vs. “I’’ (injury sprouts) and “‘S” (shaded plants) comparison. The data labeled “‘T” are identical in these separate com- parisons. Bifurcation ratios (Whitney, 1976) were computed for ramets. This is the average number of daughter branches per branch, using a branch-ordering scheme similar to that used in stream ordering (Whitney, 1976). Branch angles were measured at the base of each branch. Due to distal, adaxial bending of the branch, however, effective branch spread was less than that predicted from basal branch angles. Hence, effective branch angle (in degrees) was computed from branch length and the spread of the branch tip away from the axis of the next lower branch order. Since Aralia spinosa is deciduous, leaves could only be measured on the most recent year’s growth. A random-numbers table was used to select one leaf from each of the leafy shoots present. This leaf was characterized in terms of the annual-increment numbering system described above; leaf length and width (outer points of the outline of the compound leaf) and number of leaflets were recorded. Fifty trunk and fifty branch leaves were sampled. A subsample ANNUAL INCREMENT NUMBERING SYSTEM KEY TO ANNUAL INCREMENT NUMBERS (AL) th AI from base 3rd Al from base st a of plant of first order branch second order branch Al - Annual Increment number 1976,etc. - Year of growth INFL - Position of inflorescence T - Trunk B - Branch 7 - Terminal bud position or scars re T 1977 198] AI 2 1976 1980 a 198 Al | Al | Al ; A ~ fp RAMET AGE: 6yr 2yr lyr Figure |. Annual increment (‘AI’) numbering system used to model growth and branching of Aralia spinosa. VSONIdS VITVUV ALIHM [p861 SOP JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 a -) nH re) (@) 1 ee iN) a NODES (0°) EXTENSION GROWTH (cm) (e) $ : (0) L 1 1 l L 1 1 L r 1 9) 2 l2 ANNUAL INCREMENT NUMBER N= 96 79 87 65 32 45 40 47 37 «232 | Heceeennresseeeemesee | ee DIAM.= 1.8cem 12cm SE= .4 SE =1.0 Ficure 2. Mean length of increment (extension growth) and mean number of leaves (nodes) borne per year in Aralia spinosa as function of annual increment number. Mean diameters of 3 lowermost and 3 uppermost annual increments shown beneath figure; standard errors indicated above and below means of leaflets was used to establish a regression between leaflet blade area and dry weight. Diameters of the most recent year’s growth (one-year-old shoots) and of the basal increment of the ramet (annual increment no. 1) were measured. A subsample of ramets was harvested for determination of biomass of wood, leaves (divided into leaflet blades and rachides), inflorescences, and infructes- cences. These biomass samples were randomly selected from within the dif- ferent age classes of the ramets studied. They were oven dried at 105°C until their weight stabilized (usually after 48 hours) and then weighed. RESULTS EXTENSION GROWTH, DIAMETER, BIOMASS, AND NUMBER OF LEAVES PER SHOOT For each of the first two years, extension growth in Aralia averaged more than 75 cm; this was followed by a decline, so that by the eighth to tenth years extension growth averaged less than 25 cm annually (a 67% decline; FIGURE present; FiGuRE 2) declined, but the decline—ca. 40 percent—was less steep 1984] WHITE, ARALIA SPINOSA 407 Lor (5: | a } b os oe: ; 1.0 fe) = E L é E a. a | b. oe |.OF 2 10 me oO | é > | S a 4 oO L r) ~ > > P64 | 2 ry r) s bow fo) 5 fo) Sr 2) Pi (o) 9 5L 5 rep) : uJ oO = é 2 é Fb et 2 3 sf f a i E 1 fe) & “TB Sp F T B Sp F ol P> laa oaeee T 1 OS T | S 207 ¢. ~100r g. 2.07 = ry [e,) =~ c. 7 —_ E wn E 4 a - op) ~ q a a <> C. florida ~ ibe z P ‘ ao <> C. mas Pe Daal ce) ane an aeieae R oeareeG Week 1 qT q q q 5 T qT qT 10 T q qT y a T T T 20 T q q 25 q Week 30 March 13 May 22 July 31 Oct.9 Figure 2. Duration of bloom (lines with squares), fruit development (lines with circles), and foliage (solid lines) in Cornus spp. at Case Estates of Arnold Arboretum, March-September, 1983. (Plants still in foliage, dashed line.) When dissected and examined under the light microscope, the buds of C. mas and C. florida that were externally visible in late August were found to be flower buds; the identity—vegetative or reproductive—of those of C. sericea could not yet be discerned. These buds of Cornus species develop up to a point during the summer, remain dormant through the winter, and expand (or com- plete development) with the advent of favorable conditions the following spring to carry out their respective functions the second year. INSECT VISITORS The most commonly observed insect visitors on the species of Cornus studied were the honey bee (4pis mellifera) and the bumble bee (Bombus sp.). A third type of bee, Andrena sp., also visited Cornus mas and particularly C. sericea. Unlike the honey bee and the bumble bee, which appeared to forage mostly on pollen of these species and remained only a few seconds on each flower, individuals of Andrena sp. spent 20-40 seconds on each flower and possibly fed on the nectar. The fly species Epalpus signifer and Pollenia rudis were also seen on C. mas, but they appeared to be disinterested in either nectar or pollen and were probably casual visitors rather than foragers on this species. For insect visitors to dogwood flowers other than those reported here, see Knuth (1898), Robertson (1928), LaBerge and Ribble (1972), and Maier and Waldbauer (1979a, 1979b). 1984] GUNATILLEKE & GUNATILLEKE, CORNUS 425 BREEDING EXPERIMENTS Results of the pollination experiments are summarized in TABLE 2. In all three species of Cornus studied, fruit set in emasculated or nonemasculated, bagged flowers (la and 2a, respectively), as well as emasculated or nonemas- culated, selfed and bagged flowers (1b and 2b, respectively), was extremely low (< 3%). By contrast, plants that were cross-pollinated (1c and 2c) yielded a fruit set of 11-44 percent, depending on the species. Fruit set in C. sericea was higher than that in the other two species in both cross- and open-pollination experiments (see TABLE 2). In open-pollinated plants (1d and 2d) nonemas- culated flowers set more fruit than emasculated ones in both C. mas and C. florida but an almost equal percentage in C. sericea. However, in cross-polli- nated flowers of all three species, the emasculated flowers yielded a higher fruit set than the nonemasculated ones. In all but one instance (C. mas, 2c), cross- pollinated and bagged flowers had a higher percentage of fruit set than open- pollinated ones. In both self- and cross-pollinated flowers of all three species, pollen grains germinated on the stigmatic surface and grew through the stylar tissue. Growth was slower in the selfed flowers than in those that were crossed, however. DISCUSSION The different inflorescence types of Cornus mas, C. florida, and C. sericea appear to be related to the phenology of leaf production and to adaptations for pollination success. In C. mas flowering precedes leaf emergence. Conse- quently, the small, short-peduncled umbels are fully exposed to visiting insects. In C. florida the foliage begins to appear after flowering has commenced but before it is completed. The enlarged, showy bracts of the inflorescence heads in this species, together with their relatively long peduncles, are probably its adaptations for effective pollination. By contrast, in C. sericea leaves are formed well in advance of flower production. Here the compound cyme has not only a much larger diameter and many more flowers per inflorescence than the other two species, but also a relatively long peduncle that lifts the pollination unit well above the leaves. Although Cornus sericea came into flower last of the three species studied, it completed its fruiting cycle much sooner than the others. One of the factors contributing to its rapid fruit set could be the early growth of foliage prior to flowering and fruiting, which possibly enables the species to provide adequate food resources to the developing fruit soon after fertilization. In C. mas leaves emerged only several weeks after young fruits had formed. An intermediate situation appears to prevail in C. florida. These observations also give us some insight, little though it may be, into the resource partitioning in these species. It could be speculated that in C. mas and C. florida the previous year’s photosynthetic products provide the resources necessary for the current year’s floral and foliar expansion, whereas in C. sericea the previous year’s resources appear to suffice only for the current year’s foliage production. In this species resources necessary for flowering presumably come 426 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 from the current year’s photosynthetic products. In all three species resources for fruit development may be provided by the current year’s growth. These differences in resource partitioning may also be related to growth habit in the species studied; C. mas and C. florida are both small trees, while C. sericea is Results of controlled pollination experiments performed in this study imply several conclusions. First, the three species examined are self-incompatible, and out-crossing is obligate. Self-sterility in dogwoods, including Cornus mas, has also been previously reported (D’Amato, 1947; Hummel et a/., 1982; Ohta, 1971). The very low percentage of fruit set in self-pollinated flowers is probably due to contamination during floral manipulation. Second, the low fruit set in emasculated and bagged flowers, which again may be due to contamination, provides indirect evidence for the absence of agamospermy or apomixis. Fur- ther, because fruit set in nonemasculated and bagged—as well as selfed and bagged — flowers 1s ca. 3 percent or less, we infer that self pollen on the stigma does not stimulate or trigger agamospermy in these species. Third, microscopic observations of pollen-tube germination in stylar tissue of selfed flowers suggest that the self-incompatibility barrier in these species operates beyond the stig- matic surface. It also rules out nonviability or germination deficiency of selfed pollen as an explanation for lack of self-fertilization. The observation that ovaries (or young fruits?) of self-pollinated flowers remain a long time on the parent plant before they are ultimately shed suggests the operation of a post- zygotic barrier to self-incompatibility. However, only embryological studies will confirm whether this incompatibility is pre- or postzygotic. Fourth, the greater percentage of fruit set in emasculated and cross-pollinated flowers as compared to nonemasculated and crossed ones suggests that in nonemasculated flowers self pollen competes for space on the stigma and possibly thus con- tributes to lower fruit set. Fifth, the low fruit set in open-pollinated plants of C. florida is attributable to the absence of other genetically different individuals of this species within a radius of about 60 m of the two experimental trees, whose origin may be traced back to grafts. Sixth, although C. mas has been introduced into North America from central and southern Europe, its repro- ductive success—judged by fruit and seed set in open-pollinated flowers— indicates that local pollinators are quite effective in cross-pollinating this species. The extent to which wind contributes to pollination was not investigated in this study. ACKNOWLEDGMENTS We are indebted to the Harvard Forest of Harvard University for the Charles Bullard fellowships awarded to us, without which this study would not have been possible; to the Arnold Arboretum of Harvard University and the De- partment of Biology of the University of Massachusetts, Boston, for facilities to carry out this study; to M. Thayer for identifying the insects; to the staff of the Case Estates and the Dana Greenhouse of the Arnold Arboretum for their assistance and cooperation during field and laboratory work; and to P. S Ashton, K. S. Bawa, and P. B. Tomlinson for their advice and enthusiastic discussions during this study and for their critical comments during preparation 1984] GUNATILLEKE & GUNATILLEKE, CORNUS 427 of the manuscript. Thanks are also due to R. H. Eyde for permitting us to consult his unpublished manuscript on the Cornaceae. LITERATURE CITED Bawa, K. S. 1974. Breeding systems of tree species of a lowland tropical community. Evolution 28: 85-92. Cuan, H. T. 1977. Reproductive biology of some Malaysian dipterocarps. 107 pp. Unpubl. Ph.D. thesis, University of Aberdeen, Scotland. 1981. Reproductive biology of some Malaysian dipterocarps, III. Breeding systems. aren Forester 44: 28-36. CouLTER, J. M W. H. Evans. 1890. A revision of North American Cornaceae. Bot. Gaz. (Crawfordsville) 15: 30-38, 86-97. D’ ape F. 1947. Fenomeni di auto-incompatibilita nel Corniolo (Cornus mas L.)? o Giorn. Bot. Ital. n.s. 54: 365-367. eee ‘'H. 1932. Cytological studies of Cornus. J. Arnold Arbor. 13: 410-415. Fercuson, I. K. 1966. The Cornaceae in the southeastern United States. J. Arnold Arbor. 47: 106-116. FosBerG, F. R. 1942. Cornus sericea L. (C. stolonifera Michx.). Bull. Torrey Bot. Club 69: 583-589. Go ps att, P. 1978. A contribution to cytology in Cornales. Ann. Missouri Bot. Gard. 5: 650-655. Hummel, R. L., P. D. AscHer, & H. M. Pettetr. 1982. Genetic control of self- incompatibility i in red-osier dogwood. J. Heredity 73: 308, 309. KnutTu, P. 1898. Handbuch der Bliitenbiologie. Band 2, Teil 1. 697 pp. W. Engelmann, Leipzig. LaBerce, W. E., & D. W. Rissie. 1972. A revision of the bees of the genus Andrena of the Western Hemisphere. Part V. Trans. Amer. Entomol. Soc. 98: 271-357. Maier, C. T., & G. P. WALDBAUER. 1979a. Dual mate-seeking strategies in male syrphid flies ies Syrphidae). Ann. Entomol. Soc. Amer. 72: 54-61. —. 1979b. Diurnal so patterns of flower flies (Diptera: Syrphidae) in an eae: sand area. Ibid. 237 Martin, F. 1959. Staining and ieee pollen tubes in the style by means of fluo- rescence. Stain Technol. 34: 125-128. METCALFE, C. R., & L. CHALK. 1950. Sages Anatomy of the dicotyledons. Vol. 2. Pp. 735- 741, Clarendon Press, OntTa, Y. 1971. eee in Cane florida and C. kousa. Rep. Kihara Inst. Biol. Res. 22: REHDER, A. 1940. sical of cultivated trees and aes ime in North America. ed. 2. xxx + 996 pp. The Macmillan Company, New Ropertson, C. 1928. Flowers and insects: lists of aioe of four a and fifty- three flowers. 221 pp. Published by the author, Carlinville, Illino Sass, J. E. 1958. Botanical microtechnique. ed. 3. 228 pp. Iowa an College Press, Ames, Iowa. Seymour, F.C. 1969. The flora of New England. xvi + 596 pp. Charles E. Tuttle Co., tland, Vermont. WILKINSON, A. M. 1944. Floral anatomy of some species of Cornus. Bull. Torrey Bot. Club 71: 276-301. Witson, J. S. 1965. Variation of three taxonomic complexes of the genus Cornus in eastern United States. Trans. Kansas Acad. 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The following Congress Symposia are being organized: Symbiosis in Evolution Conservation of Tropical Ecosystems Biogeographic Evolution of the Malay Archipelago Adaptational Aspects of Physiological Processes Co-evolution in Ecosystems and the Red Queen Hypothesis Angiosperm Origins and the Biological Consequences The Measurement of Rates of Evolution Molecular Biology and Evolutionary Theory Co-evolution and Systematics Molecules vs. Morphology in Phylogeny: Conflict or Compromise? Random and Directed Events in Evolution Biochemical Innovation in Microbial Communities There will also be Special Interest Symposia on other topics, as well as sessions for contributed papers, films, and poster papers. For further information write to: Professor Barry Cox ICSEB Congress Office 130 Queen’s Road Brighton, Sussex BN1 3WE England Journal of the Arnold Arboretum July, 1984 CONTENTS OF VOLUME 65, NUMBER 3 A Taxonomic Revision of the American Species of Agarista (Erica- ceae). Wo I 2b tte ee cee eueancaruaeenre cawaneapeeonacs 255 The Tribes of Cruciferae (Brassicaceae) in the Southeastern United States. IHSAN A. AL-SHEHBAZ ....... a ee ee aay eee en are” 343 Inflorescence Architecture and Evolution in the Fagaceae. MOnERT B; RAUL AND ERNST C. ABEE 4554<255466092dc0580440 6% 375 The Architecture of Devil’s Walking Stick, Aralia spinosa (Araliace- ae). PETER 9. WHITE: 3.6%. 0 00526220 $35080%4 Radda bibdaddines das 403 Some Observations on the Reproductive Biology of Three Species of Cornus (Cornaceae). C. V. S. GUNATILLEKE AND I. A. U. N. GUNATILLEKE ........... 419 Volume 65, Number 2, including pages 149-254, was issued April 16, 1984. JOURNAL ot te ARNOLD ARBORETUM HARVARD UNIVERSITY VOLUME 65 NUMBER 4 ISSN 0004-2625 Journal of the Arnold Arboretum Published quarterly in January, April, July, and October by the Arnold Arboretum, Harvard University. 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JOURNAL OF THE ARNOLD ARBORETUM VOLUME 65 OcTOBER 1984 NUMBER 4 SYSTEMATICS OF CENTRAL AMERICAN HELICONIA (HELICONIACEAE) WITH PENDENT INFLORESCENCES! W. JOHN KREss THE FIRST botanical description of Heliconia was by Plumier in his Nova Plantarum Americanarum Genera (1703). He provided a short generic de- scription of the genus Bihai and polynomials for three taxa: Bihai amplissimis Soliis, florum vasculis coccineis, Bihai amplissimis feliis, florum vasculis sub- nigris; and Bihai amplissimis foliis, florum vasculis variegatis. In his Species Plantarum (1753), Linnaeus included these three taxa in a single species, Musa bihai, retaining Plumier’s exact diagnoses and placing the “variegatis” variety first. Miller (1754, 1768) and Adanson (1763) considered these plants generically distinct from other species of Musa and used the generic name Bihai. In Mantissa Plantarum (1771), Linnaeus also segregated M. bihai into its own genus, Heliconia L. He provided a generic description of Heliconia and gave a short diagnosis of a species, H. bihai, with red cincinnal bracts and yellow flowers. Kuntze (1891; substituting the variant spelling Bihaia) and later Griggs (1904, 1915) recognized the earlier generic name and transferred all species of Heliconia known to them into Bihai Adanson. However, at the International Botanical Congress held in Vienna in 1905, Heliconia was rein- stated as a nomen conservandum (Farr et al., 1979). Around the turn of the century, a number of workers attempted revisions or summaries of the genus. Petersen (1890) and Kuntze (1891) each listed 25 species. Baker (1893), the first to provide an infrageneric classification of the genus, supplied descriptions and keys for 29 species. In 1900 Schumann sum- marized the genus for Engler’s Das Pflanzenreich but added no new species and simply translated abbreviated versions of Baker’s earlier descriptions into Latin. Griggs was one of the most knowledgeable students of Heliconia, due rt of a thesis (Kress, 1981b) submitted to the Department of Botany, Duke University, in partial fulfillment of the requirements for the Ph.D. degree. © President and Fellows of Harvard College, 1984. Journal of the Arnold Arboretum 65: 429-532. October, 1984. 430 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 in part to his study of plants in the field. However, he never produced a thorough revision, although he did publish several papers on the genus (1903, 1904, 1915). The last of these papers recognized 48 Neotropical species and contained an infrageneric classification that was more detailed than the one proposed by Baker in 1893. Since the treatment by Griggs in 1915, there has been no major revision of the genus in its entirety. During the last 65 years, many new species have been described and several regional floristic treatments have added to our knowledge of the variation and diversity within Heliconia. Lane began work on the genus at Harvard University in the late 1940’s and annotated many herbarium spec- imens with names of new species and varieties, but unfortunately he never published any of these taxa. Recently, Andersson (1981) has provided a modern revision of the species related to H. bihai (sect. HELICONIA) that is based pri- marily on herbarium material and has solved many of the taxonomic and nomenclatural problems of that group. Floristic treatments are available for the heliconias of Guatemala (Standley & Steyermark, 1952), Costa Rica (Stand- ley, 1937), Panama (Woodson & Schery, 1945), Middle America (Smith, 1968), Venezuela (Aristeguieta, 1961), Peru (Macbride, 1936), and the Old World (Green, 1969), but most of the studies are now out of date and incomplete. Recent workers in Costa Rica (Daniels & Stiles, 1979), Nicaragua (Smith, 1977, 1980), Panama (Kress, 1981a), Colombia (Abalo & Morales, 1982, 1983b), Ecuador (Dodson & Gentry, 1978; Abalo & Morales, 1983a), and Brazil (Bar- reiros, 1971, 1976, 1978; Emygdio, 1975, 1976; Santos, 1978) have described many new species. The work of Daniels and Stiles (1979) in particular is an excellent example of the importance of field work in understanding taxonomic delimitations within Heliconia. INFRAGENERIC CLASSIFICATION Several infrageneric classifications of Heliconia have been proposed (see TABLE 1), each one more complex and composed of more hierarchical ranks than its predecessor. Early authors based their groups solely on the shape of the cincinnal bracts. Later workers also used plant size, leaf orientation, and inflorescence habit and structure to devi detailed d supposedly more natural—classifications. Kuntze (1891) published the first infrageneric taxon above the rank of species. The two species contained within sect. Taeniostrobus Kuntze—Heliconia im- bricata and H. mariae—are characterized by imbricate cincinnal bracts. Baker (1893, p. 190) divided the genus into two subgenera, Platychlamys Baker and Stenochlamys Baker, the former containing all species with ‘“‘ovate-acuminate, deeply boat-shaped”’ cincinnal bracts; the latter, species characterized by bracts that are “lanceolate-acuminate, shallowly boat-shaped.”’ Schumann (1900) fol- lowed Baker’s classification except that he placed Platychlamys in synonymy under Taeniostrobus and altered the ranks of these taxa from subgenera to sections. Griggs’s superior knowledge of the genus was reflected in his more complete and less artificial classification. He recognized (1903, p. 644) that infrageneric 1984] KRESS, HELICONIA 43] TABLE 1. History of the infrageneric classification of Heliconia. BAKER (1893) SCHUMANN (1900) Griccs (1903) Gricas (1915)* Subg. Platychlamys Sect. Taeniostrobus Subg. Taeniostrobus Subg. Taeniostrobus Baker Kuntze Kuntze Kuntze Episcopales Griggs Subg. Stenochlamys — Sect. Stenochlamys Subg. Platychlamys Imbricatae Griggs Baker Baker Baker Champneianae Griggs Pendulae Gri Subg. Stenochlamys Subg. Stenochlamys Baker Baker Distantes Griggs Cannoideae Griggs *Under the generic name Bihai. groups based upon a single Se avai bract shape, inadequately ex- pressed species relationships: “‘It i t to be denied that the shape of the branch-bracts [cincinnal bracts] is in a ee way correlated with the rela- tionships of the species, but it is only an accidental parallelism without much physiological importance, for there are many exceptions—species quite similar in all respects except that the branch-bracts are sufficiently different to place them in different subgenera... .’’ In a diagnostic key he included three sub- genera, Stenochlamys, Platychlamys, and Taeniostrobus, defined by plant habit and cincinnal-bract orientation. Griggs later (1915) replaced this classification with one consisting of two subgenera (7aeniostrobus and Stenochlamys) and six subordinate taxa of un- specified rank. All Species known at that time were included in his treatment. Plant habit and height rientation, and distance between adjacent cincinnal bracts were characters used to recognize subgeneric groups. However, the actual phylogenetic relationships within Heliconia are much more complex than is represented in Griggs’s final classification. Some question exists as to whether Griggs formally proposed the six sub- ordinate taxa in his 1915 publication because he did not specify their rank. The International Code of Botanical Nomenclature (Stafleu, 1978) specifies that a new name published before 1953 does not require clear indication of rank. In addition, although no formal descriptions or diagnoses were provided, the taxa were sufficiently differentiated in the key provided by Griggs in the same publication. The names for the infrageneric taxa comply with all the other ee of the Code for valid publication at that time. Griggs’s infrage- eric names must therefore be recognized as validly published but without facet rank. In the present treatment, I do not propose any new classification of Heliconia. Rather, I have purposefully chosen to study in depth a group of species—those with pendent inflorescences—that has previously been classified as a natural assemblage. Where possible, these species are compared with others not in- cluded in this taxonomic group. In addition to broad morphological compar- 432 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 isons, I have used information on breeding systems, results from artificial and natural interspecific hybridization, and analyses of pollen structure to assess the phylogenetic status of the assemblage. The information provided here will hopefully establish a foundation upon which a more natural classification of the entire genus can eventually be constructed. Griggs (1903, 1915) was the first to suggest that species of Heliconia with pendent inflorescences are closely interrelated, and he included most of these species in his unranked Pendulae. However, by placing several species that he described as having pendent inflorescences (H. punicea, H. mariae, and H. curtispatha) into a group characterized by erect inflorescences (Jmbricatae), he indicated that inflorescence habit was not necessarily an indication of close relationship. The present revision includes all Mexican and Central American taxa of Heliconia with pendent i no known species with pendent inflorescences in the Antilles). The nineteen species and eight varieties involved fall into three of Griggs’s lower groups, /mbricatae, Pendulae, and Distantes. One object of this study was to test the hypothesis that the pendent inflorescence habit is a uniquely derived state in the genus that characterizes a monophyletic group of species. Ideally, all pendent species, including those from South Amer- ica, should have been incorporated into the study. Unfortunately, a total re- vision was not logistically possible due to the large number of species, the wide geographic distribution, and the extensive field work needed to investigate the group adequately. For this reason the conclusions reached here concerning the Pendulae can only underestimate the complexity of this infrageneric group. Because of this complexity and our lack of knowledge, it is still premature to assign any rank to Griggs’s Pendulae. A final statement on the phylogenetic classification of Heliconia must await a treatment of the entire genus. An extension of the present revision of the Central American species with pendent inflorescences to include the South American species (approximately 50) is planned. THE HELICONIACEAE The proper classification of the order Zingiberales has been debated since 1880, when Bentham and Hooker first established four tribes within the Sci- tamineae. Subsequent authors have subdivided and altered the rank of each of the original four tribes (see TABLE 2). Most current workers recognize eight families within the order. Heliconia has always been allied with the Musa complex that includes Or- chidantha N. E. Br., Musa L., Ensete Horan., Strelitzia Banks, Ravenala Adan- son, and Phenakospermum Endl. These genera are recognized as distinct from other members of the Zingiberales by their usually arborescent habit and their flowers with five (or six) pollen-bearing stamens. Lane (1955) presented con- vincing evidence for segregating Orchidantha into a monotypic family, the Lowiaceae (proposed by Hutchinson, 1934). He retained the remaining genera in a single subdivided family, the Musaceae. Subdivision of the family into several ranks, Lane believed, best shows the relationships of the genera. He TABLE 2. Systems of classification.* ass \o fore) Nakai (1941 & TOMLINSON (1962) Cronauist (1981) BENTHAM & HOOKER ENGLER & PRANTL ENGLER — 1902, ENGLER & PRANTL UTCHINSON GREN & CLIFFORD (1880 (1889) 1904, (1930) ae 1959) (1982) Order: Scitamineae No rank Order: Ce Order: Scitamineae Order: Scitamineae Order: Zingiberales Tribes Families Families Families Families Families Zingibereae Zingiberaceae Zingiberaceae Zingiberaceae Zingiberaceae Zingiberaceae Subfamilies Subfamilies Tribes Zingiberoideae Zingiberoideae Zingibereae Hedychieae is Globbeae tH Costoideae Costoideae Costeae Costaceae fs Maranteae Marantaceae Marantaceae Marantaceae Marantaceae Marantaceae an Canneae Cannaceae Cannaceae Cannaceae Cannaceae Cannaceae se Museae Musaceae Musaceae Musaceae Musaceae Musa O (Musa, Ravenala, Tribes Subfamilies Subfamilies (Musa) ee Pus oO Strelitzia, Museae usoideae usoideae Z Heliconia) ; Sa (Musa) > Ravenala, Strelitzia) eer Strelitzioideae Strelitziaceae Strelitziaceae ribes Oe (Strelitzia Grains ee. avena Ravenala, (Strelitzia, (Strelitzi en Phenakospermum) avenala) Sears ) Heliconia) Heliconieae Heliconieae Heliconieae Heliconiaceae (Heliconia) (Heliconia) (Heliconia) (Heliconia) Lowioideae Lowioideae Lowiaceae Lowiaceae aN ww ww *After Tomlinson (1962). 434 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 argued that Heliconia is the most divergent genus, more closely allied to Musa than to the other three genera. After extensive anatomical studies of the order, Tomlinson (1962) concluded that all eight divisions of the Zingiberales exhibit about the same degree of morphological and anatomical differentiation, and that each should be regarded as a family. He therefore followed the suggestion of Nakai (1941) and recognized the Heliconiaceae (including only Heliconia) and the Strelitziaceae (comprising Strelitzia, Ravenala, and Phenakospermum) as distinct from the Musaceae (containing Musa and Ensete). Recent studies on pollen of the Zingiberales (Kress et al., 1978; Stone et al., 1979, 1981) support Tomlinson’s morphological and anatomical evidence for the recog- nition of the Heliconiaceae, and his classification is accepted here. Other recent authors (Stebbins, 1974; Takhtajan, 1980; Cronquist, 1981; Dahlgren & Clif- ford, 1982) have also adopted this classification. MORPHOLOGY The distinctive morphology of members of the genus Heliconia merits a short descriptive summary. The following descriptions are taken from obser- vations of living plants and in some cases are not readily applicable to dred herbarium specimens. Although not all character states discussed (e.g., shoot habit, leaf venation, pedicel twisting) are applicable to species with pendent inflorescences, they are nonetheless included to facilitate comparison with other species and to aid workers unfamiliar with the genus. LiFE ForRM The genus comprises medium to large erect herbs often with extensive sym- podial rhizomatous growth (Bell & Tomlinson, 1980). The patterns of pro- duction of | rhizomes, thizomatous branches, and erect shoots result in varying capacities fo d differ among species. An investigation of i ee: Gaiation in rhizome morphology and branching pattern may yield information of taxonomic value. Each erect shoot is composed of a pseudostem and leaves, and it is often—although not always—terminated by an inflorescence (FIGURE 1). The pseudostem is made up of an axis covered by overlapping sheathing leaf bases, can be up to several meters in length, and has various colors and textures. In some species (e.g., Heliconia platystachys) the pseudostem has a distinctive white, waxy bloom LEAVES Leaf arrangement is distichous. If the leaves are oriented vertically and have long petioles, the plants have the aspect ofa banana plant and are called “‘Musa- like’ (FicureE 1, A). If the leaves are more or less horizontally positioned and the blades are short petiolate or essentially sessile on the pseudostem, the plants have the aspect of a ginger plant and are called “ Zingiber-like” (FiGuRE 1, C). These terms correspond to the somewhat confusing terms ““musoid”’ and “‘can- noid,” respectively, of other authors (Schumann, 1900; Griggs, 1915; Smith, 1984] KRESS, HELICONIA 435 Ficure 1. Schematic representation of the 3 types of shoot organization in Heliconia: A, Musa-like; B, Canna-like; C, Zingiber-like. 1968; Daniels & Stiles, 1979; Andersson, 1981; Abalo & Morales, 1982, 1983a). Some species may have short or medium-length petioles with blades that are held obliquely, and they indeed have a shoot organization resembling that in species of Canna. Such plants are called ‘‘Canna-like” (Ficure 1, B). The leaves of a single shoot in Zingiber-like species usually lie in a single plane. This same planar configuration is characteristic of some Musa-like species (€.g., Heliconia maculata, H. trichocarpa) whose leaves often fan out from stream banks or from the slopes of small embankments. Lateral veins run perpendicular or oblique to the central midrib (Tomlinson, 1959). Abaxial venation, although usually green and obscure, is sometimes distinctive in color (e.g., in Heliconia reticulata), and the midrib usually differs from the lamina in color and texture both adaxially and abaxially. The lamina is usually green, but in some species (e.g., Heliconia ramonensis, H. secunda var. viridiflora) it is often tinted maroon or red abaxially, especially along the margin. In most species it is elliptic-oblong, coriaceous, entire, and glabrous adaxially and abaxially. Notable exceptions are oblanceolate blades (in the trichocarpa group), those that split into narrow lateral segments with age (e.g., in H. magnifica), and those having a thick, white, waxy bloom abax- ially (e.g., H. curtispatha, H. collinsiana var. collinsiana). The apex is acute to acuminate, and the base is nearly always unequal, with one side extending farther along the petiole. In most cases the base is obtuse to truncate, but it can be cordate, or even attenuate along the petiole (e.g., in H. maculata). 436 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 INFLORESCENCES Inflorescences are almost always terminal on erect, leafy shoots, but in a few species (e.g., Heliconia metallica) they may arise on a basal leafless shoot. When terminal, the inflorescence may have an erect or pendent orientation with respect to the leafy shoot from which it emerges (FiGuRE 1). In some taxa (e.g., H. secunda, H. sessilis, and some hybrids between erect and pendent species) the inflorescence is carried in an intermediate, nodding posture. The peduncle (the part of the stem between the terminal leaf sheath and the basal cincinnal bract; Figure 2, A) may have various colors and textures. The in- florescence is made up of modified leaflike structures called cincinnal bracts (‘“‘branch-bracts” or “‘spathes”’ of previous authors), the rachis connecting ad- jacent cincinnal bracts, and a cincinnus of flowers within each bract. Mea- surements of inflorescence length do not include the peduncle. The rachis (FiGurE 2, B) may differ from the cincinnal bracts in color and texture and is either straight or flexuose (zigzag). Rachis-internode length is measured between two adjacent cincinnal bracts (FiGurRE 2, C). The cincinnal bracts are distichous or are spirally arranged due to twisting of the rachis. In some species (e.g., 1. xanthovillosa) the rachis is only slightly twisted, making the cincinnal bracts subspirally arranged. Each bract is oriented at an angle of from 0 to 180° to the axis of the inflorescence (FiGuRE 2, F). The cincinnal bract closest to the peduncle is the basal bract, is often sterile, and may be elongated and leaflike (FiGuRE 2, D). The most conspicuous feature of a fertile plant is the colorful inflorescence. The cincinnal bracts are usually bright red and/or yellow but are sometimes green (e.g., H. talamancana) or even pink (e.g., H. colgantea). In some species (e.g., H. stilesii, H. maculata) the bracts are glabrous or essentially so, while in others (e.g., H. magnifica, H. xanthovillosa) the entire inflorescence may be covered by long, woolly hairs. Inflorescence and flower parts may be glabrous, puberulous, tomentose, velutinous, villous, or woolly. The term scur- fy is used for any exfoliating vestiture that is easily removed by slight abrasion. The color and texture of the cincinnal bracts generally differ on the inside and outside surfaces. Because the cincinnal bracts decrease in size toward the apex of the inflorescence, those situated in the middle portion of the inflorescence (FIGURE 2, E) are best for comparative measurements. Bract length is measured from the rachis to the distal tip (FicuRE 3, Aa). The width is not measured across the open top since this changes with age, but from margin to margin as if the bract were spread out flat (FiGuRE 3, Ab). Mean length/width quotients (1/w) are indicative of general bract shape. Bracts with quotients greater than 2 have long-acuminate apices; those with quotients less than 1.3 have short- acuminate or acute apices. The margins of the cincinnal bracts may be straight, revolute, or involute near the rachis. FLORAL BRACTS Each flower of the cincinnus is subtended by an individual floral bract (FIGURE 2, G). Often there are several basal floral bracts that do not directly subtend any individual flower. The floral bracts of some species are opaque and cori- aceous, persisting through fruit development to protect maturing ovaries, while 1984] KRESS, HELICONIA 437 Ficure 2. Structure and measurements of inflorescences of Heliconia: A, peduncle: B, rachis; C, rachis internode length; D, basal cincinnal bract (sterile, with elongated leaflike extension); E, middle cincinnal bract; F, cincinnal bract angle with axis of inflo- rescence (e.g., 80°); G, floral bract those of other species are filmy or translucent and quickly decompose after anthesis. The floral bracts are variously colored and are textured abaxially; the adaxial surface is always glabrous. FLOWERS Although most floral cl teri y preserved in pressed and dried plants, they can be quite helpful and even eee when the flowers are fresh (or, in some cases, rehydrated). The flowers are hermaphroditic. Each cincinnus contains several to many (up to 50) flowers. The pedicel (FiGuRE 3, Bg) is 438 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Structure and measurements of cincinnal bracts and flowers of Heliconia. URE 3. A, cincinnal bract (Aa, length; Ab, width; Ac, flower protruding at anthesis). B, flower (Ba, perianth length; Bb, perianth width; Be, perianth angle (e.g., 90°); Bd, apex of fused sepal reflexed; Be, apex of fused sepal not reflexed; Bf, ovary; Bg, pedicel). C, perianth, outer surface showing 2 sepals fused to partially spread-open corolla tube. sta- minodes: D, abaxial view; E, position relative to style, lateral view. F, style and stigma. usually short and obscured by the floral bracts, but it can be exposed and distinctive in some species (e.g., Heliconia collinsiana, H. trichocarpa). The perianth is made up of two whorls united at the base that show varying degrees of fusion within and between whorls. At anthesis the single adaxial sepal be- comes free from the other perianth members (FiGurE 3, Ac) and allows legit- imate pollinators to enter the floral tube. The apices of the abaxial sepals are free from the corolla tube and may be reflexed (Ficure 3, Bd) or not (FIGURE 1984] KRESS, HELICONIA 439 A —E 4. Staminode shape in Heliconia (not to scale): A, acuminate; B, cuspidate; C, aie D, bidentate; E, lobed. 3, Be). The corolla tube is split along the margins of the two adaxial petals. The partially fused corolla tube is adnate to the two fused abaxial sepals (FIGURE 3, C) and lies opposite the free sepal. The free sepal opens above in erect inflorescences and below in pendent ones. In some species with erect inflores- cences (e.g., H. imbricata), the pedicel is twisted, causing the flower to be resupinate and therefore to open in a fashion opposite to that described above. The perianth is usually yellow, varying from pale yellow to white at the base and to deep yellow apically. In some taxa the flowers are green (e.g., H. secunda var. viridiflora) or pinkish red (e.g., H. mariae). The sepals can be glabrous or pubescent, and the free sepal is sometimes velutinous or even woolly (e.g., pogonantha group). The length and curvature of the perianth tube reflects the length and curvature of the bill of the pollinating hummingbird. The length of the perianth is measured from its articulation with the ovary to the tip of the free adaxial sepal, following any curvature of the tube (FiGurE 3, Ba); the width at its widest diameter (FiGuRE 3, Bb). Following the terminology of Andersson (1981), the perianth tube may be slightly curved (e.g., H. collinsiana), parabolic (e.g., H. stilesii, FiGurE 3, Bc), or sigmoid (e.g., H. trichocarpa). Five stamens are fertile and produce viable pollen. The anthers, borne at the ends of long filaments that traverse the length of the perianth, are either connivent inside the corolla or flared outside of the perianth apex, and they dehisce longitudinally. The sixth stamen is replaced by a staminode located opposite the free sepal (FiGure 3, D, E). This modified stamen does not produce pollen but may function in some species as a guide leading the hummingbird’s tongue to the floral nectaries situated at the base of the style. The staminode ranges in length from 5 to 20 mm, and the apex may be acuminate, cuspidate, apiculate, dentate, or lobed (FiGure 4). The ovary (FiGure 3, Bf) is inferior and three-locular. Each locule contains a single, basally attached ovule. The hypanthium is white, yellow, or green, and although it is usually glabrous, in some species (e.g., Heliconia trichocarpa, H. maculata) it is puberulous to pubescent. The style is much elongated, runs 440 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 the entire length of the perianth, and may be geniculate near the stigma (FIGURE 3, F). The lobed stigma is surrounded by the five fertile anthers at the apex of the perianth. Stigma morphology (degree of lobing and papilla development) appears to vary among species and may prove diagnostic in future studies. FRuITS Previous authors have described the fruits as berries, capsules, and “schizo- carpic berries.” An early study by Humphrey (1896) on seed development in the Zingiberales demonstrated that the mature fruit of Heliconia is a drupe with a stony endocarp enclosing each of the true seeds. This interpretation has recently been reconfirmed (B. Kirchoff, pers. comm.) and is accepted here. The outer pericarp is fleshy, and at maturity the surface layer becomes blue (Neo- tropical species) or red to orange (Paleotropical species). The fruits are very attractive to birds that disperse the seeds (Skutch, 1933; Stiles, 1979). Each drupe contains from one to three pyrenes. Each seed is surrounded by an exceptionally hard, roughened endocarpic envelope. Unlike the seeds of many other members of the order, those of Heliconia do not have an aril. The embryo is poorly differentiated at the time of seed maturity (Gatin, 1908), which may account for the delayed germination of Heliconia seeds often encountered by horticulturists. A Note TO COLLECTORS If an entire inflorescence cannot conveniently be collected, several represen- tative sections (including cincinnal bracts from the base and apex) and the peduncle should be pressed. It is helpful if one cincinnal bract is cut open to reveal the flowers and floral bracts. A portion of the leaf showing the base of the blade and a portion showing the apex should also be collected and pressed. Several vegetative and reproductive characters are of taxonomic importance but are usually not preserved after pressing and drying. Collectors should record the states of these characters from living specimens in the field: Pseudostem: Color and vestiture Leaf: Configuration Musa-, Canna-, or Zingiber-like Orientation spiral or planar Color of lamina (adaxial and abaxial) Color of midrib (adaxial and abaxial) Inflorescence: Position terminal or arising from a basal shoot Habit erect or pendent Color of peduncle Color of rachis Color of cincinnal bract (inside and outside) Arrangement of cincinnal bracts distichous or spirally arranged 1984] KRESS, HELICONIA 441 Flower: Orientation resupinate or not resupinate Color of perianth and ova Curvature of apices of fused sepals reflexed or straight Orientation of anthers flared outside perianth apex or connivent inside apex Shape of staminode POLLEN The pollen of Heliconia and most of its allies in the Zingiberales (excluding Zingiberaceae and Costaceae) is characterized by a sporoderm significantly different in structure from that of most angiosperms. The normally elaborate protective exine is reduced to a few spinules and a thin (0.08 wm) connective layer covering the surface. The intine, contrastingly, is much thickened (8 um) and structurally complex (Erdtman, 1966; Kress et a/., 1978). Radially striate channels perforate the intine of the distal hemisphere, while a more homoge- neous but stratified intine characterizes the proximal hemisphere. Recently, the ultrastructure (Kress et al., 1978), cytochemistry (Kress & Stone, 1982), and ontogeny (Stone et al/., 1979) of Heliconia pollen have been described in detail. A systematic comparison of pollen of 27 Central American species of Heliconia has been published elsewhere (Kress & Stone, 1983). The results of that study that are pertinent to the present taxonomic revision are briefly summarized here. The pollen of heliconias with pendent inflorescences is 46-65 um in polar diameter and 54-86 um in equatorial diameter. The distal radius is 24-56 um; the proximal radius 8-29 um. The grains may be euoblate, suboblate, or oblate- spheroidal, and isopolar, subisopolar, or heteropolar. In addition to variation in size, the shape and sculpturing of each hemisphere, which are generally different in a single grain, also vary among species. The distal hemisphere is usually convex (but may be subconvex or planar) and sometimes is truncate or shows constrictions parallel to the equatorial plane. The proximal hemi- sphere may be convex, subconvex, planar, or subconcave. Although the entire wall of the distal hemisphere resembles the germination aperture of more conventional angiosperm pollen, the grains appear inaperturate. However, the pollen tube invariably emerges from a central region at the distal pole. This central region is the functional germination “‘aperture”’ and is often structurally distinct. It may protrude from the pole and be elongated, or it may be rather flattened and partially sunken into the distal surface. Sculpturing of the reduced exine ranges from psilate to granulose to verrucose to spinulose or minutely spinulose. In most species the pattern is different on the two hemispheres. Statistical analysis of both quantitative and qualitative pollen characters support the recognition of four groups among the 19 species with pendent inflorescences studied: the pogonantha group, including Heliconia pogonantha, H. magnifica, H. ramonensis, H. danielsiana, H. xanthovillosa, and H. mariae; the curtispatha group, comprising H. curtispatha and H. stilesii; the trichocarpa 442 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 group, with H. trichocarpa, H. colgantea, H. necrobracteata, H. maculata, and H. talamancana, and the nutans group, containing H. nutans, H. collinsiana, H. secunda, H. platystachys, and H. marginata. The polar diameter, the degree of grain polarity, the shape of each hemisphere, the shape of the germination aperture, and the sculpturing of the proximal hemisphere are of particular taxonomic value in delimiting these groups, which are also characterized by other morphological traits (see section on phylogeny). The pollen of H. sessilis, however, shares some character states with each of the four groups and so is not easily placed in any of them. Pollen of the pogonantha type (FiGurE 5, A) is large (60-65 x 81-86 um), euoblate to oblate-spheroidal, and heteropolar, with a convex-truncate, spi- nulose distal hemisphere and a planar, psilate proximal hemisphere. The ger- mination aperture is distinct, flattened, and somewhat sunken. Pollen of the curtispatha type (Ficure 5, B) is medium sized (53-57 x 66- 70 um), suboblate, and isopolar, with a convex to subconvex, spinulose distal hemisphere and a convex, minutely spinulose proximal hemisphere. The ger- mination aperture is distinct and usually sunken. Pollen of the trichocarpa type (FIGURE 5, C) is medium sized (49-55 x 67- 79 um), euoblate, and isopolar, with a subconvex to planar, verrucose to spi- nulose distal hemisphere and a convex, granulose to minutely spinulose prox- imal hemisphere. The germination aperture is indistinct except for an occa- sional slight depression in the center of the planar distal face. Pollen of the nutans type (Ficure 5, D) is relatively small (43-56 x 54-76 um), euoblate to oblate-spheroidal, and subisopolar, with a convex, often con- stricted, spinulose distal hemisphere and a subconvex to planar, psilate to verrucose proximal hemisphere. The germination aperture is distinctly pro- truding and sometimes quite elongate. CYTOLOGY The cytology of Heliconia is poorly known. Of the 14 species for which chromosome numbers have been reported (Bisson ef a/., 1968; Mahanty, 1970), fewer than half have legitimate names and several are not identified to species. In none of the studies was any voucher specimen designated, and it is unlikely that the majority of specimens were identified correctly. In addition, since only one of the species listed (H. nutans, if identified correctly) has a pendent inflorescence, the chromosome counts reported are of little help in the present study HABITAT AND GEOGRAPHIC DISTRIBUTION Members of the genus Heliconia are distributed primarily throughout the New World tropics from the Tropic of Cancer in Central Mexico to the Tropic of Capricorn in South America. Most species inhabit moist or wet regions, but some are found in seasonally dry areas. Although heliconias attain their most luxuriant vegetative growth in the humid lowland tropics at elevations below 500 meters, the greatest numbers of species (many locally endemic) are found 1984] KRESS, HELICONIA 443 Ficure 5. Pollen types of Heliconia with pendent inflorescences: A, pogonantha type (H. pogonantha, distal view); B, curtispatha type (H. stilesii, distal view); C, OED type = trichocarpa, distal view); D, nutans type (H. nutans, equatorial view). Scale 10 um in middle-elevation rain and cloud-forest habitats. Few species occur above 2000 meters. The most conspicuous members of the genus inhabit open sites in secondary growth along roadsides, on river banks, and in forest light gaps. With increased destruction by man of the tropical rain forest, these species readily invade and colonize the newly opened areas. Other species never attain such extensive vegetative growth and are restricted to the more shaded habitats of the primary forest. These latter species are often locally endemic and are fast becoming extinct as destruction of the forest accelerates Species of Heliconia with pendent inflorescences are found in Mexico, Central merica, South America, and Melanesia (the Solomon Islands). None is known from the West Indies. The taxa treated in this revision are distributed from 444 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Michoacan in southern Mexico to Darién in Panama. Most species occurring north of Costa Rica are found on the wet Atlantic slopes and coastal plains. In Costa Rica and some parts of Panama, the Pacific slopes have sufficient rainfall to support a number of species of Heliconia. In these countries several pairs of closely related species (e.g., H. curtispatha/H. stilesii, H. trichocarpa/ H. colgantea, H. pogonantha/H. danielsiana) are allopatrically distributed on Opposite: sides of the central cordillera. No differences in habitat or geographic t between species with pendent and erect inflorescences, and species of the two inflorescence habits are often sympatric. A curious disjunct group of heliconias is found in the Old World tropics. These heliconias are distributed from Samoa westward to the central Indo- nesian island of Sulawesi. These plants undoubtedly belong in the genus, even though a separate generic name, Heliconiopsis, has been suggested (Miquel, 1859). Green (1969) included all of the Old World taxa in a single species, Heliconia indica Lam. A taxonomic treatment currently in preparation (Kress, unpubl.) recognizes eight species occurring in this area, some locally endemic to specific islands or land masses. Two of these species (both unpublished) have pendent inflorescences. POLLINATION AND REPRODUCTIVE BIOLOGY Several original reports and review papers describing the reproductive bi- ology of Heliconia have recently appeared (Stiles, 1975, 1979; Kress, 1981b, 1983a, 1983b, in press). Only the taxonomically important aspects will be summarized here; see the cited papers for details. In the Neotropics hummingbirds are the exclusive pollinators of Heliconia. Species-specific relationships between birds and plants are rare. Any Heliconia, however, can be categorized as being visited primarily either by traplining hermit hummingbirds or by territorial nonhermit hummingbirds (Linhart, 1973; Stiles, 1975). Physiological self-incompatibility is uncommon in the genus; most species that have been tested so far are self-compatible (Kress, 1983a). Plant/bird morphological specialization, hummingbird foraging behavior and habitat, and phenological differences may all serve as isolating mechanisms that prevent pollen exchange between species (Stiles, 1975, 1978; Kress, 1983b, in press). Of all the mechanisms restricting interspecific gene flow in Heliconia, physiological regulation of foreign pollen germination and pollen-tube growth at the stigmatic surface and in the style is the most important (Kress, 1983b). I have found that in the species tested, pollen of any one species was inhibited at the stigma, within the stylar tissue, or within the ovary of most other species, and that the site of inhibition depended on the species combination and the direction of the cross. Cross-compatibility was uncommon, and hybrid seed was obtained from very few hybrid combinations. Natural hybridization is also rare (Kress, 1981b, 1983b, in press), especially considering the large number of sympatric taxa in the genus. The few hybrids between species with pendent rau rte known from the wild are discussed in the section on natural hybridizati The ee of the interspecific crossability barriers is significant from a 1984] KRESS, HELICONIA 445 taxonomic standpoint. In my study (Kress, 1983b) at Las Cruces Tropical Botanical Garden in Costa Rica, nine species with pendent inflorescences were crossed in nearly all reciprocal combinations. Each species was categorized by the ability of the style to accept foreign pollen tubes and the ability of the pollen to grow in foreign styles. Species were then grouped together according to the main type of crossability barrier that characterized the pistil and pollen of each (see TABLE 3). For example, in Heliconia pogonantha, H. danielsiana, and H. colgantea foreign pollen is inhibited at the stigmatic surface, and pollen of these species is inhibited on the stigma of most foreign species. In contrast, although foreign pollen is nearly always inhibited at the stigma in H. collinsiana and H. nutans, the pollen of these species breaks the stigmatic barrier of most other species, and pollen-tube growth is arrested in the style. In the case of H. mariae, H. stilesii, and H. curtispatha, foreign pollen is stopped at the stigma and/or style, and their own pollen does not penetrate foreign stigmas. Heliconia trichocarpa has the least specific pistil- and pollen-inhibition barriers of the nine species tested. When compared to the groups of species recognized by their vegetative, floral, and pollen characters (see section on phylogeny), the groups defined by crossability barriers are nearly the same, even though the placement of some species (e.g., H. colgantea and H. mariae) 1s inconsistent. Presence of specific crossability barriers appears to reflect the phylogenetic history of at least some species of Heliconia. PHYLOGENY One goal of taxonomic investigations is to reconstruct phylogenetic rela- tionships. Phylogenetic systematics (sensu Hennig, 1966), or cladistics, pro- vides a logical and repeatable method for formulating hypotheses of phylogeny and for devising biological classifications (Bremer & Wanntorp, 1978; Eldredge & Cracraft, 1980; Wiley, 1981). Cladistics requires that a classification rec- ognize only monophyletic groups (groups that include all and only the descen- dents of a common ancestor) and hence be isomorphic with the hypothetical genealogy of the taxa. Monophyletic groups can only be recognized on the basis of shared uniquely derived character states (synapomorphies); shared primitive or ancestral character states (symplesiomorphies) can only define paraphyletic groups (those that include some but not all of the descendents of an ancestor). The most compelling argument for adoption of phylogenetic systematics and for using cladistic hypotheses as the basis for classification is that a common genealogy is the sole factor that unites all organisms. A cladistic analysis of the 19 Central American species of Heliconia with pendent inflorescences (hereafter referred to as ““‘pendent species’’) was under- taken in an attempt to answer two questions: 1) What are the genealogical relationships of the taxa? 2) Is the group of species with pendent inflorescences monophyletic (i.e., is the pendent inflorescence habit shared by these species a synapomorphy that defines a monophyletic group)? The first question can be answered by constructing a cladogram for the taxa, as outlined below. A definitive answer to the second question is more difficult to obtain using only the 19 species included in the revision because these species represent a subset 446 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 TABLE 3. Main sites of pistil and pollen inhibition in species of Heliconia crossed at Las Cruces Tropical Botanical Garden.* MAIN SITE OF INHIBITION SPECIES Pistil Pollen H. colgantea H. danielsiana Stigma Stigma H. pogonantha H. collinsiana ; : pinane Stigma Stigma/style H. curtispatha HT. mari Stigma/style Stigma Hi. stilesii H. trichocarpa Stigma/style Stigma/style *Table from Kress, 1983b, © Annals of Botany Company, 1983; reprinted with permission. of all heliconias. To provide at least a tentative answer to this question, five species with erect inflorescences (hereafter referred to as “erect species’’) were added to the phylogenetic analysis of the pendent species. If either inflorescence habit is uniquely derived in Heliconia, then it should define a monophyletic group of species, all of which have that inflorescence type. MetTHops, TAXA, AND CHARACTERS The basic methodology of phylogenetic systematics includes definition and delimitation of monophyletic taxa as operational units for the analysis; careful and accurate assignment of character states to taxa based upon sound hypoth- eses of homology; formulation of hypotheses of character-state polarity using outgroup comparison; construction of a cladogram of the taxa; careful mappin of character-state changes on the cladogram and reassessment of the original hypotheses of monophyly, homology, and polarity; and, ifappropriate, devising a hierarchical classification based on the monophyletic groups defined by the cladogram. The results of phylogenetic systematic studies are only meaningful if the taxa under investigation are monophyletic (Hennig, 1966; Eldredge & Cracraft, 1980; Arnold, 1981; Wiley, 1981). It is normally assumed that species are monophyletic. In the present analysis each of the species used as a unit taxon for construction of the cladograms has a suite of apomorphies that suggests it is a monophyletic lineage (see individual species descriptions). The monophyly of the entire group is a central question of the investigation. The presence of a pendent inflorescence in all of the species suggests that the 19 Central Amer- ican species as a group may be a monophyletic lineage. If the pendent inflo- rescence is a uniquely derived character in Heliconia, the exclusion of all South American pendent species automatically makes it likely that the group 1s para- phyletic. Nonetheless, the limitations that result from not including all mem- bers of a monophyletic group in a phylogenetic analysis do not invalidate the 1984] KRESS, HELICONIA 447 cladogram as the best hypothesis of relationships of the included taxa. The first cladogram presented here, which includes only the Central American species with pendent inflorescences, therefore provides a testable hypothesis of the genealogical relationships of these taxa. To test the hypothesis of monophyly of the pendent species, a second clado- gram incorporating five additional species with erect inflorescences was con- structed. These particular five species were chosen for several reasons. Com- parative data on character-state distribution (especially information on pollen characters) were available for the species. The five species are representatives of both of Griggs’s (1915) subgenera (Taeniostrobus and Stenochlamys) and three of his five unranked supraspecific groups that contain erect species (Jm- bricatae, Distantes, and Cannoideae). The majority of the pendent species fall into his sixth supraspecific taxon, Pendulae. Heliconia imbricata and H. wag- neriana, together with H. mariae and H. curtispatha (with pendent inflores- cences), were classified in Griggs’s Jmbricatae, a group of species with imbricate or overlapping cincinnal bracts. Heliconia wagneriana has most recently been included by Andersson (1981) in sect. HELICONIA, which contains members of Griggs’s Imbricatae and Champneianae. Heliconia latispatha and H. tortuosa were members of Griggs’s Distantes, a group of species with erect inflorescences and a Musa-like shoot habit. The latter species is very similar to H. secunda (with a pendent inflorescence) and therefore suggests some affinity between pendent and erect species (Stiles, 1979). Heliconia mathiasiae, although not yet described when Griggs developed his classification, would have been in- cluded in his Cannoideae because of its “‘cannoid” (here called Zingiber-like) shoot habit. The morphological diversity of the five erect species and their representation of most of Griggs’s supraspecific groups of Heliconia insures that a reasonable sample of different species was chosen to test the hypothesis of the monophyletic status of the pendent species. If synapomorphies could easily be recognized and distinguished from sim- ilarity due to homoplasy or plesiomorphy, delimitation of monophyletic groups would be simple and unambiguous. The fact that accurate identification of synapomorphies is difficult requires careful and explicit character analysis, specially with regard to comparison of homologous structures. The only phy- logenetically useful concept of homology is genealogical: two taxa share a feature ecause it was present in their common ancestor (Eldredge & Cracraft, 1980). The hypotheses of homology required to construct a cladogram, however, must initially be based on indirect sources of evidence (e.g., similarity in position, anatomy, or development). The resulting cladogram, representing the largest nested set of mutually consistent, hierarchically correlated characters, will then provide a test of these hypotheses of homology. Characters not consistent with the cladogram (those requiring multiple independent origins or reversals) are homoplasious and suggest that the initial hypotheses of homology for these characters were incorrect and in need of reevaluation. Character states assigned to each of the species (see TABLE 4) for the 30 characters used in the analysis were determined for at least three individuals per population for one to several populations throughout the geographic range of each taxon (TABLE 5). Discrete, nonoverlapping states for characters with 448 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 potentially continuous distributions (e.g., overall height, inflorescence size) were determined by the use of bar graphs. Whenever possible, intraspecifically variable characters were eliminated from the analysis. In some cases in which a species was variable for a given character, the state present in its closest relatives was chosen as the primitive state for the variable species. These characters were then reevaluated after the cladogram was constructed. For example, the four varieties of Heliconia pogonantha vary in the presence of hairs on the inflorescence. The velutinous perianth present in all varieties of this species is also found in other closely related heliconias that have velutinous to woolly inflorescences. A hairy inflorescence was therefore chosen as the primitive state in the variable H. pogonantha. Outgroup comparison, the most logically justifiable method for determining character-state directionality (Lundberg, 1972; Eldredge & Cracraft, 1980; Ste- vens, 1980; Watrous & Wheeler, 1981), was used to polarize the characters in Heliconia (see TABLE 6). By general consensus (TABLE 2; Baker, 1893; Lane, 1955; Tomlinson, 1962; Dahlgren & Clifford, 1982), members of the “banana- like” families of the Zingiberales, especially the Musaceae and Strelitziaceae, are considered to be the closest relatives of Heliconia. Several shared features of floral morphology unite the Heliconiaceae most closely with the Musaceae, and a recent attempt to construct a phylogeny of the entire order (Dahlgren & Rasmussen, 1983) recognizes these two families as sister groups. For the cla- distic analyses of Heliconia, the two genera of the Musaceae sensu stricto, Musa and Ensete, were chosen as the outgroup. Information on character states in the Musaceae was taken from Lane (1955), Tomlinson (1962), Erdtman (1966), Argent (1976), Dahlgren and Clifford (1982), and Kress and Stone (unpubl.). Character states universally present in this family were scored as primitive in Heliconia. Characters not present in the outgroup (e.g., hummingbird polli- nation), or present with more than one state (e.g., inflorescence habit), were not used. Multistate characters were coded as either unidirectional (0 > | > 2; characters 8, 12, 17-21, 24, 25) or bidirectional (0 — 1 — 2; characters 7, 10, 11, 13) according to indirect evidence for character-state homologies (see TABLE 6). For clarity, auttapomorphies for single species of Heliconia and char- acter states present in the genus but not in the species with pendent inflores- cences were also eliminated. In total, the 30 characters used in the analysis contained 44 character-state changes or evolutionary steps (TABLE 6). The cladogram was constructed using Farris’s Wagner ’78 computer program. The program is based on a Wagner tree algorithm (Kluge & Farris, 1969; Farris, 1970) that produces the most parsimonious tree without placing any restrictions on homoplasious evolution. One problem with the program is that 1t may produce cladograms of different lengths depending on the order in which the taxa appear in the data file (Coombs et a/., 1981). However, in these analyses the same cladograms were produced regardless of the order in which the taxa were submitted. RESULTS CLADOGRAMS. The cladogram of the 19 species with pendent inflorescences (cladogram I; FiGureE 6), including a hypothetical ancestor having all hypoth- 1984] TasLe 4. Taxon by character matrix for 24 species of Heliconia and 30 characters. Character* Taxon cLcGtt COL KRESS, HELICONIA HAHA OCH COnFCCONnNO COHN CON AAA MOON NAAAN TH HOCOCOAOANN eA On AA CCH AAA AOC N eo0o0on COW OH OOCOCCO oooconoooon ooco°o HoOoOCOCHnWOOCOCCCCCHO oooocooooc°oon ooo OOn nF OOCOOCOTFOCOnAAOCOOd 0 010 0 0 2 *See Table 6 for character states and polarity. THypothetical ancestor haviag all primitive character states. 449 TFor species abbreviations see Table 5. + 1 450 JOURNAL OF THE ARNOLD ARBORETUM TABLE 5. [voL. 65 Field collections of Heliconia species from which ser pai state data were taken for phylogenetic analyses and descriptions of ta TAXON* COLLECTION DATAT H. colgantea Daniels & Stiles (CLG) H. collinsiana a (COL) var. collinsia var. velutina Kress H. curtispatha Petersen (CUR) H. danielsiana Kress (DAN) ** HT, imbricata (Kuntze) Baker (IMB) ** 7. latispatha Bentham (LAT) H. maculata Kress (MAC) H. magnifica Kress (MAG) H. marginata (Griggs) Pittier (MRG) H. mariae J. D. Hooker (MAR) **H, mathiasiae Daniels & Stiles (MAT) H. necrobracteata Kress (NEC) H. nutans Woodson (NUT) H. platystachys Baker (PLT) H. pogonantha Cuf. (POG) var. pogonantha var. holerythra Daniels & Stiles var. pubescens Daniels & Stiles var. veraguasensis Kress H. ramonensis Daniels & Stiles (RAM) var. ramonensis var. glabra Kress var. lanuginosa Kress var. xanthotricha Kress Costa Rica: Puntarenas, 77-878 Guatemala: Alta Verapaz, 77-747 Esquintla, 77-753 Nicaragua: Managua, 77-758 Guatemala: Quezaltenango, 77-756 nama: Panama, 77-864 ee Rica: Sa 77-821 é, 79-1097 Costa Rica: eee 79-1101 Panama: Panama, 77-850, 80-1246 Costa Rica: Puntarenas, 77-804, 79-1095 Guatemala: Petén, 77-743 ta Verapaz, 77-745 Costa Rica: slates 77-771 ve 79-1104 , 77-794 ees soy Zelaya 77-761 nama: Canal Zone, 77-855 ie a Heredia, 79-1105 Panama: Coclé, 77-842, 80-1158 Panama: Chiriqui, 77-822, 79-]088, -1089a, 79-108 9b, 82-1344, §2-1345, 82-1355, 82-1368 Veraguas, 82-1410 Costa Rica: Puntarenas, 77-803 Panama: Canal Zone, 77-860 Costa Rica: Heredia, 77-778, 102 Nicaragua: Zelaya. 77-760 Costa Rica: Cartago/Limé6n, 77-788 Panama: Veraguas, 77-825 Costa Rica: Alajuela, 77-764 Panama: Coclé, 8 9 Panama: Chiriqui, 83-1600, 83-1602 Panama: Coclé, 77-840, 80-1161 1984] KRESS, HELICONIA 451 TABLE 5 (continued). TAXON* COLLECTION DATAT H. secunda R. R. Smith (SEC) var. secunda Costa Rica: Heredia, 77-773, 77-770 var. viridiflora Daniels & Stiles Costa Rica: Guanacaste, 80-1216 H. sessilis Kress (SES) Panama: Colon, 77-871, 80-1241 H. stilesii Kress (STI) Costa Rica: Puntarenas, 79-1096 San José, 77-799 H. talamancana Daniels & Stiles (TAL) Costa Rica: Limon, 77-791 **H tortuosa Griggs (TOR) Costa Rica: Puntarenas, 78-944 H. trichocarpa Daniels & Stiles (TRI) Costa Rica: Alajuela, 77-767, 77-804 Cartago, 77-787 Heredia, 79-1/09 Panama: Veraguas, 77-832 ** 1H wagneriana Petersen (WAG) Panama: Coldén, 80-1244 H. xanthovillosa Kress (XAN) Panama: Panama, 80-1233, 83-1565 *AYT the author. Species with erect inflorescences are indicated by a double asterisk (**). +All collections were made by the author and are deposited in the Duke University Herbarium (pUKE). For complete locality information, see “Specimens Examined” under the appropriate species description. for species names used in phylogenetic analysis are given in parentheses following =) esized plesiomorphic character states, had a total length of 84 evolutionary steps. Forty steps were homoplasious. The index of consistency (Kluge & Farris, 1969), which is the total length of the cladogram minus the homoplasies divided by the total length, was .524. This index permits comparisons of the degree of homoplasy or parsimony between cladograms constructed from different char- acter sets or containing different taxa. Character-state changes for each of the 30 characters are indicated on the cladogram. The second cladogram (cladogram II; Ficure 7), which includes the 19 pendent and the five erect species, had a total length of 96 steps. Fifty-two steps were homoplasious, and the index of consistency was .458. The same basic topology of the four monophyletic groups of pendent species present in cladogram I was maintained in cladogram II. The five erect species do not form a monophyletic lineage. Three of the erect species are intercalated within pendent lineages: Heliconia mathiasiae in the trichocarpa group, and H. /ati- spatha and H. tortuosa in the nutans group. The remaining two erect species, H. wagneriana and H. imbricata, constitute two single-species monophyletic groups that do not share a most recent common ancestor. CHARACTER ANALYSIS. Twenty of the 30 characters used in the construction of cladogram I had at least one uniquely derived state (characters 2-5, 7-13, 20, 22-25, 27-30). Some characters, such as overall height (character 1) and pollen- grain shape (characters 19, 21), were homoplasious and not phylogenetically 452 JOURNAL OF THE ARNOLD ARBORETUM TABLE 6. Thirty characters used in the phylogenetic analyses. CHARACTER CHARACTER STATES (values) —_ nN ww BSS ws On ~ co — — SO . Overall shoot height . Pseudostem surface . Pseudostem vestiture . Leaf-blade shape . Shoot aspect . Inflorescence size (number of bracts) . Rachis vestiture . Cincinnal-bract length/width quotient . Cincinnal-bract margins . Cincinnal-bract outer vestiture . Cincinnal-bract inner vestiture . Flower shape . Vestiture at perianth apex . Anther orientation . Staminode length . Ovary vestiture . Pollen polar diameter . Pollen equatorial diameter . Overall pollen shape (P/E) not glaucous (0)* glaucous (1) absent (0)* present (1) oblong-elliptic (0)* oblanceolate (1 ect (o} horizontal/ebligue (1) essentially absent (1)* short hairs (0)t long hairs 0) <1.4 (0)* 1.5-2.0 (1)t 0 (2) straight (0)* involute at base (1) s eaeuriae! absent (1)* s (0) essentially absent (1)* short hairs (0) long hairs (2) essentially absent (1)* short hairs (0) long an 0) connivent inside perianth apex (0)* flared outside perianth apex (1)t absent (0)* short hairs (1) 260 um (0)* 49-59 um (1)t <48 um (2) > 80 (0)* 65-79 um (1)t (2) oblate- Saar (=.88) (0)* 1) suboblate (.7 euoblate (<. i a [voL. 65 1984] KRESS, HELICONIA 453 TABLE 6 (continued ). CHARACTER CHARACTER STATES (values) NO oO . Pollen polarity (DR/PR) isopolar (<2) (0)* subisopolar (2-5) (1) heteropolar (> 5) (2) 21. Pollen distal-hemisphere shape I convex (0)* subconvex (1)t planar (2) 22. Pollen distal-hemisphere shape II not truncate (0)* truncate (1) 23. Pollen distal-hemisphere shape III not constricted (0)* constricted (1) 24. Pollen distal-hemisphere sculpturing psilate (0)* granulose—minutely spinulose (1)f spinulose (2) 25. Pollen proximal-hemisphere shape convex (0)* 26. Pollen proximal-hemisphere sculpturing Cal (0)* anulose—minutely spinulose (1)t 27. Pollen germination aperture I Saeed (O)* distinct (1) 28. Pollen germination aperture II not protruding (0)* protruding (1) 29. Pollen germination aperture III not sunken (0)* sunken (1) 30. Pollen germination aperture IV not flattened (0)* flattened (1) *Character state present in outgroup and originally coded as primitive for Heliconia. tCharacter state present in most recent common ancestor of species in cladogram I. informative. Several other features of the pollen grains (characters 22-25, 27- 30) proved quite useful in delimiting monophyletic lineages (also see Kress & Stone, 1983). Long hairs on inflorescences and flowers (characters 7, 10, 11, 13) are uniquely derived in one monophyletic group (see below), whereas short hairs appear to have evolved independently in several ene and are therefore not homologous in all heliconias in which they occu The large number of character-state changes eee 6-8, 12, 14, 17-19, 21, 24, 26) occurring between the hypothetical ancestor and the extant heli- conias in cladogram I suggests that the most recent common ancestor of these pendent heliconias did not have some of the originally assigned plesiomorphic states present in the outgroup. These eleven characters were recoded so that the primitive state is the one present not in the outgroup but in the most recent common ancestor (see TABLE 6). In addition, states of multistate characters present in the outgroup but not in the pendent heliconias were eliminated from the transformation series of characters 12, 21, and 24. The result is a more 454 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 TRI GRP NUT GRP j CUR GRP Ar POG GRP Ll Jt ees | NEC MAC TAL CLG TRI MRG SEC NUT PLT COL CUR STI SESMAR POG RAM DAN MAG XAN | i 1070 170 I 21:2 15:0 1:0 13: oc A 1 1:0 152] 1 9:0 ‘ 16:1 23:0 12:2 17:0 13:0 18:1 7:2 : Ww NMR adores CoHo Re 1 7 t 16 28 7:0 1 21:0 1 3 1 6:0 29:1 l 7:0 > l } 1 rhe 241 26 i ANC Ficure 6. Cladogram I. Character-state changes in each lineage given as character number followed by state value. All species have pendent inflorescences. Species groups indicated by horizontal bars above taxa names. (For character states and species abbre- viations, see TABLE 6.) parsimonious cladogram (i.e., with fewer homoplasies) without any alteration in the branching sequence of cladogram I. This revised cladogram (not shown) has eleven fewer homoplasies, six additional synapomorphies (characters 6, 8, 14, 17, 18, 26), and a higher index of consistency (.586). Three characters (7, 19, 21) showed no significant change in homoplasy after the recoding. Although recoding the characters in this fashion does not affect the classification, it does provide new hypotheses on the attributes of the most recent common ancestor of the included pendent species and requires a reevaluation of those character states that are uniquely derived in the various monophyletic lineages. These character-state changes are incorporated into the discussion of monophyletic MONOPHYLETIC GROUPS. Four primary monophyletic groups or lineages of pendent species are present in both cladograms I and II (Ficures 6, 7): the trichocarpa group (Heliconia necrobracteata, H. maculata, H. talamancana, H. colgantea, and H. trichocarpa), the nutans group (H. marginata, H. secunda, H. nutans, H. platystachys, and H. collinsiana); the curtispatha group (/. (‘9 ATaV] 99s ‘suONeIAaiqge salads pue soje]s 19}0eIeYD 10.4) i SOWWBU BXB] DAOQE sieq [e}UOZLIOY Aq paleorpul sdnoss saisads "SQOUDISIIOYU!I JUSpUad JABY S1OYI0 |]B “YSLIoIse uv AQ Pa1VoIpUI SsadUadSaIOYUT eta YIM Saidadg ‘anjea aje1s AQ PaMO][O} JoquNU Ja}OBIeYO se UDATS BBROUT] YOO UI SadURYS d1RIS-1O}IeIeYD “[] wWesopeg “1 awNo1y ONV lib2 T6L 1:8 T:2t lal 1:9 142 1:2 2:2 1:02 Teil pm 16 < A es2l 0 Q — — ea) 1:92 2:12 - ve at a 2:61 2:8 ce 1:8 Us jaa) vez sn Z a £3 | gues Utz 1:91 EE REISS I O01 OF 8T UB aS 2:62 O24 ORT oFeT OFSL OFT ase tT os SS WVY 50d YVAN S3S ILS YNOGWI 10D Ld YOL LAN 03S LV 1 SYN OVM = DTO OVW WL DAN ee inl tle [eas cae ee ae Os ee a oe ae? Swe | Oe dy) 50d dy) HNd dy LINN dy ly 456 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 curtispatha and H. stilesii), and the pogonantha group (H. sessilis, H. mariae, . pogonantha, H. ramonensis, H. danielsiana, H. magnifica, and H. xan- thovillosa). The five members of the trichocarpa group are united with each other and distinguished from other heliconias by several shared derived character states. Oblanceolate leaves that are widest toward the apex and tapering toward the base, shoots that tend to be held in an oblique or a horizontal plane, and a cincinnal bract l/w quotient greater than 2 (except in Heliconia necrobracteata) are synapomorphies of this group. Two pollen characters, granulose to minutely spinulose distal-hemisphere sculpturing and indistinct germination apertures, are unique to this group, but they are also present in the hypothetical ancestor shared by all heliconias. These characters are therefore not synapomorphies but do identify members of the group. Staminodes over | cm long are nearly unique to members of the trichocarpa group (except H. maculata) but are also present in H. collinsiana of the nutans group. Similarly, puberulous to tomen- tose ovaries are primarily found in the trichocarpa group (except H. necro- bracteata) but are also present in a single species of the nutans group, #7. platystachys. Synapomorphies of the nutans group include slightly curved flowers and pollen with protruding germination apertures. Two characters, cincinnal bracts with a l/w quotient between 1.5 and 2, and pollen grains with a subconvex proximal hemisphere, are primarily restricted to the nutans group but are also found in its common ancestor with other heliconias and are therefore not synapomorphies. The nutans group is more heterogeneous than the trichocarpa group. Although Heliconia marginata is united by various apomorphies with the other species in this group, it differs in its involute cincinnal bract margins, in its medium-size euoblate pollen grains lacking constricted distal hemi- spheres, and in several autapomorphies. Heliconia marginata may have closer relatives among South American taxa. Heliconia collinsiana and H. platysta- chys are united by the presence of a waxy pseudostem, but each has several states unique within the nutans group. Heliconia collinsiana has large shoots, long staminodes, and pollen grains without a constriction of the distal hemi- sphere, states that have evolved independently outside the nutans group. Hel- iconia platystachys has pubescent cincinnal bracts and ovaries and large pollen grains—characters unique within the nutans group but homoplasious within the genus. The remaining pendent heliconias are members of the sister curtispatha and pogonantha groups. Large inflorescences with many cincinnal bracts, a cincin- nal bract I/w quotient less than 1. 5, an cosas within the perianth apex, and pollen grains with sunken g states uniquely derived in the common ancestor of these ‘two groups. The curtispatha group has no synapomorphies that distinguish it from all other heliconias but does have several unique states that are absent in its sister pogonantha group. For example, puberulous to tomentose cincinnal bracts (inner and outer surfaces) and isopolar pollen grains with convex proximal hemispheres are not unique to the curtispatha group but distinguish it from the pogonantha group. Several unique pollen-grain features present in the pogonantha group are planar, psilate proximal hemispheres and flattened germination apertures. Two 1984] KRESS, HELICONIA 457 additional pollen-grain character states, heteropolarity and a truncate distal emisphere, are synapomorphies of the pogonantha group minus Heliconia sessilis. This latter species is only loosely united with the other species of this group and may represent a Central American species with closer phylogenetic ties to South American taxa not included in this revision. Similarly, H. mariae is the sister species to the remaining taxa of the pogonantha group, although it has several features (straight cincinnal-bract margins, parabola-shaped flow- ers, and puberulous perianth apices) that distinguish it from the other members. A velutinous to woolly vestiture on the rachis, cincinnal bracts (outer and 1 inner diameter exceeding 80 um), an additional shared derived feature of these five species, are also independently derived in H. platystachys of the nutans group. The pogonantha group as a whole is more heterogeneous than either the cur- tispatha or the trichocarpa group. EVOLUTION OF THE INFLORESCENCE HABIT. Evidence from cladogram II (FIGURE 7), which was constructed using the original hypotheses on character polarities (TABLE 6), supports the hypotheses that both pendent and erect inflorescences have most likely evolved several times in He/iconia and that pendent heliconias do not constitute a monophyletic group. The five species with erect inflores- cences do not form a separate monophyletic lineage but are incorporated into several different lineages of pendent species. Synapomorphies supporting the placement of each erect species are indicated on the cladogram. Inflorescence habit was not used as a character in the construction of clado- gram II. Because the outgroup includes species with both pendent and erect inflorescences, the inflorescence habit of the common ancestor of all heliconias is uncertain. According to cladogram IJ, if the ancestor had an erect inflores- cence, the most parsimonious arrangement of state changes of this character requires five evolutionary steps: either three independent origins of pendent inflorescences and two reversals to the erect state, or two separate origins of the pendent state and three reversals to the erect state. If the ancestor had a pendent inflorescence, then five independent origins of the erect habit are required to map the character on the cladogram most parsimoniously. Re- gardless of the inflorescence type of the common ancestor, this character is very homoplastic in Heliconia, and therefore neither state is homologous in all taxa in which it occurs DISCUSSION Before the present investigation of Heliconia, few attempts had been made to analyze the phylogenetic relationships within the entire genus or any part of it. Smith (1968), using less than ten characters, constructed a phylogenetic tree for the Middle American species of Heliconia, but provided no method- ology or rationale to support his conclusions. All of the species with pendent inflorescences were included in a single lineage that excluded all erect species. Stiles (1979, p. 151) placed 37 Costa Rican species into seven groups based 458 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 on morphology and phenology and suggested that these groups may correspond to “real evolutionary or taxonomic relationships.” He did not state any explicit method by which he formulated his groups, but his decisions have in part been confirmed by the present analyses. Stiles categorized all of the pendent species into two groups, the pogonantha group and the trichocarpa group. Stiles’s pogonantha group, corresponding essentially to the curtispatha and pogonantha groups defined here, would be monophyletic according to cladogram I. As pointed out earlier, the curtispatha group has no states uniquely derived within Heliconia, and it could therefore be treated as a sublineage within a more broadly defined pogonantha group, as was done by Stiles. However, the set of derived character states present in the curtispatha group but absent in the pogonantha group supports recognition of the separate taxonomic status of the two lineages. Stiles’s second pendent group, the frichocarpa group, contains species that are here placed in the trichocarpa and nutans groups and would therefore be paraphyletic according to cladogram I. Heliconia marginata, a somewhat aberrant member of the nutans group, was listed as incertae sedis by Stiles. He further pointed out that some species with pendent inflorescences (e.g., his trichocarpa group—especially H. secunda) are very similar to some species with erect inflorescences (e.g., his tortuosa group), suggesting that all pendent species should not be classified in a single group or even in closely related groups. This suggestion has been substantiated by cladogram II. Three of the four pendent monophyletic groups discussed in this revision contain some taxa that are not restricted to Central America (e.g., Heliconia marginata, H. platystachys, H. curtispatha, H. mariae, and H. pogonantha). In addition, each of the four lineages certainly contains one or more South American species not considered here. The heterogeneity of the nutans and pogonantha groups is probably due to the fact that some of these species (e.g., H. marginata, H. platystachys, H. sessilis, and H. mariae), which are quite different from other members of their groups, may have closer relatives in South America than in Central America. The inclusion of only Central American species requires some caution in the interpretation of the relationships of the taxa in cladograms I and II. However, incorporating pendent South American species or more species with erect in- florescences into the analysis most likely will not require major reinterpretation of the main evolutionary lineages defined here. The addition of more taxa may expand the monophyletic lineages or create new sublineages within the major groups but will not change the relationships among the taxa presently included. For example, Heliconia mathiasiae, the sister species of the trichocarpa group, has an erect inflorescence and a Zingiber-like shoot habit. If other heliconias with Zingiber-like shoots were added to the analysis, they would most certainly be placed in the same lineage with H. mathiasiae. Taken together, these Zin- giber-like species would then be the sister group of the frichocarpa group. The overall relationships of the other pendent and erect species outside this lineage would not be altered. This reasoning also applies to the inclusion of other species with erect or pendent inflorescences that would be intercalated into the cladogram as sister groups to various species without altering the overall re- lationships of the other lineages. This was demonstrated here by the incor- 1984] KRESS, HELICONIA 459 poration of the five species with erect inflorescences in a reanalysis of the pendent species. Neither the shone of the pendent species nor the cir- cumscription of the four ma tin cladogram I was greatly changed in cladogram II by ae addition of the extra taxa. With regard to evolution of inflorescence type, the relatively small number of species included in the cladistic analysis here probably underestimates the degree of homoplasy in this character. An analysis of the entire genus may suggest an even greater number of independent origins of both pendent and erect inflorescences. CONCLUSIONS The cladistic analyses support the recognition of at least four separate mono- phyletic lineages of Central American Heliconia with pendent inflorescences. The lineages are here called the trichocarpa group, the nutans group, the cur- tispatha group, and the pogonantha group. The first two groups include some species with erect inflorescences as well. These results suggest multiple inde- pendent origins of each inflorescence type and therefore argue against the in- clusion of all heliconias with pendent inflorescences in a single monophyletic lineage that excludes all species with erect inflorescences. SPECIES CONCEPTS The strong physiological isolating mechanisms that in most cases prevent fertilization between species of Heliconia (Kress, 1981b, 1983b) would seem to support the biological species concept (Mayr, 1957, 1969). The many sym- patric species of the genus that share habitats, flowering times, and pollinators, as well as the few natural hybrids that are produced, suggest severe restrictions on gene flow between species. Even closely related species (e. g., HA. trichocarpa in any species of Heliconia is lacking. In the studies on crossability barriers, only tests between species were attempted and no conclusions have been reached concerning crossability between populations of each taxon. For this reason a biological species concept for ae is dificult to accept without resorting to ad hoc hypotheses on gene Simpson (1961) and Wiley (1978, 1981) have suggested a species concept that does not rely exclusively on reproductive continuity between populations of a species but does accept it as an important component of species cohesion. The evolutionary species is “‘a lineage of ancestor-descendant populations which maintains its identity from other such lineages and which has its own evolu- tionary tendencies and historical fate” (Wiley, 1981, p. 25). A lineage consists of one or more populations that share a common history of descent not shared 460 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 by other populations. Identity is interpreted as the assemblage of morphological (and behavioral) characters that permits “recognition” between organisms. Reproductive isolation may of course be necessary for maintenance of the genetic identity of an evolutionary species. However, morphological coherence of any species, especially one that is distributed over large geographic areas, 1s more likely a consequence of genealogy than of gene flow between its members. Boundaries between species of Heliconia were defined by morphological, ecological, and geographic discontinuities. Botanists have variously advocated narrow and broad concepts when awarding species status to recognized taxo- nomic entities of Heliconia. Actually, the taxonomic rank designated by the taxonomist may be of little importance in understanding the biology of the organisms. The duty of the systematist is to document the extent of natural variation, record the levels of discontinuities, and define monophyletic groups. In the present study of Heliconia, differences in color, size, and shape of sexual reproductive structures that have been observed in the field have been given high priority in making taxonomic decisions. Assemblages of individuals shar- ing distinctive cincinnal-bract and flower characteristics that appear to affect pollinator visitation are awarded species status. These morphological charac- teristics are always correlated with geographic discontinuities. Varietal status is accepted for assemblages of individuals showing minor morphological dis- tinction and allopatric distributions. Taxonomic inflation has been avoided where possible, but not at the expense of obscuring information that 1s indis- pensible to other biological investigations. MATERIALS AND METHODS The relatively few taxonomic studies of Heliconia and the lack of any recent comprehensive revision of the genus are in part due to the difficulties of pre- paring representative herbarium specimens. Although usually quite common in the Neotropics, heliconias are poorly collected because of their large stature and their fleshy nature, and collections that are made are often unimpressive and uninformative. In preparing the present revision, I have emphasized ex- tensive work with living plants in the field; I have seen all of the taxa described here and have studied them in their natural habitats. Morphological data for the taxonomic descriptions, diagnostic keys, and phylogenetic analysis have been taken from living plants in situ, and voucher specimens are deposited in the Duke University Herbarium (DUKE). From one (for the rarer taxa) to seven populations (for the more widely distributed taxa) have been sampled for each taxon (see TABLE 5). Three or more a different individuals have been included in each population samp In addition to the field studies, numerous specimens from various herbaria (A, BM, CR, DUKE, F, GB, GH, K, M, MO, NY, PMA, S, SCZ, U, UC, US, and w) have been examined. Information from these specinicns has bees used to supplement field observations on morphology, habitat, geographic distribution, and phe- nology. 1984] KRESS, HELICONIA 461 TAXONOMIC TREATMENT Heliconia L. Mant. Pl. 2: 147. 1771, nom. cons. Type species: Heliconia bihai L.) L Bihai Miller, Gard. Dict. abr. ed. 4. Vol. | [alph. order]. 1754, nom. rejic. Bihai Adanson, Fam. Pl. Pt. 2. 67. 1763, nom. rejic. Heliconiopsis Miq. Fl. Indiae Batavae 3: 590. 1855. Type species: Heliconiopsis am- boinensis Miq. Bihaia Kuntze, Rev. Gen. Pl. 2: 684. 1891, orth. var. Medium to large rhizomatous herbs with Musa-, Canna-, and Zingiber-like habits, forming clones with erect, leafy shoots in groups of few (1 or 2) to many (>50). Pseudostem composed of overlapping sheathing leaf bases of various colors and textures. Leaves usually large, either distichous with all blades lying in a single plane or appearing spirally arranged; petiole short (Zingiber-like), medium length (Canna-like), or long (Musa-like); blade with the base usually unequal on 2 sides of midrib, cordate to attenuate, the upper surface usually green, the lower surface green to maroon. Inflorescence terminal on leafy or leafless shoot, erect, pendent, or nodding, consisting of brightly colored pe- duncle, rachis, and few (3 to 5) to many (>30) cincinnal bracts; cincinnal bracts distichous or spirally arranged, each subtending cincinnus of few (3) to many (>40) flowers. Flowers hermaphroditic, each subtended by opaque or membranous, variously colored floral bract, persistent or decomposing after anthesis; pedicel short (5 mm) to long (2 cm); perianth consisting of 2 whorls united at base with various degrees of fusion within and between whorls, the calyx with 2 free abaxial sepals adnate to corolla except at apex, and | nearly free adaxial sepal usually reflexed at anthesis, the corolla with 3 petals, connate except for free margins opposite free sepal; pollen-bearing stamens 5, the fil- aments long, linear, attached to base of perianth tube, the anthers 4-loculate, linear, situated at apex of perianth at anthesis, dehiscence longitudinal; pollen large, pseudoinaperturate, oblate to spheroidal, heteropolar with reduced sur- face ornamentation; staminode |, opposite free sepal, varying in size and shape; ovary inferior, 3-celled, the ovules solitary, erect, the style filiform, straight, or geniculate, the stigma capitate or lobed. Fruits 1- to 3-seeded drupes, usually blue. Seeds surrounded by stony, roughened endocarp, exarillate; embryo straight; endosperm copious. KeEY TO CENTRAL AMERICAN SPECIES OF HELICONIA WITH PENDENT INFLORESCENCES The following key was constructed from living plants and hence is most useful in the field. If sufficient label data are supplied on herbarium sheets, however, the key can be used to identify dried specimens as well. Some char- acters (e.g., pollen-grain features) will be of limited diagnostic value to most workers, but they are included for completeness. The key is based primarily on the phylogenetic tree (FIGURE 6) and is therefore a “‘natural”’ one that reflects the phylogenetic history of the taxa. Members of each of the four species groups 462 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 (trichocarpa, nutans, curtispatha, and pogonantha groups) are placed together in adjacent entries. A. parr emai small (< 20 cincinnal bracts); cincinnal ‘bract Uw > 1.4; anthers flared outside perianth apex; pollen g distinctly protruding but not nen B. Shoots oriented horizontally or obliquely; leaves oblanceolate, widest toward apex and tapering toward base; flowers sigmoid; pollen germination aperture indistinct. C. Cincinnal bract l/w < 2; ovaries glabrous. ........... 1. H. necrobracteata. C. Cincinnal bract 1/w > 2; ovaries puberulous to villous Rachis and floral bracts usually glabrous, perianth green or yellow-green, slightly sigmoid; staminode apex acuminate or apicula . Shoots < 3 min height; pseudostem pale green with dark brown spots; cincinnal bracts red, becoming green distally; perianth glabrous, yel- low, becoming green distally. .................... . H. maculata. E. Shoots usually > 3 m in height; ase es ee cincinnal bracts mostly green; perianth puberulous, mostly green. ................. sis Hawhssehiel ease aunt nen sianae se ae alae eee nee 4. Hi. talamancana. Rachis and floral bracts deat to tomentose; perianth yellow, strong- ly sigmoid; staminode apex F. Cincinnal bracts bright oe often becoming green toward apex, pu- berulous; floral bracts stiff, conspicuous; perianth pieriaee ae te eet Spa tayjey. eddie neous Tae eee Hi. colgantea. F. Cincinnal bracts red, often becoming maroon ne apex, glabrous; floral bracts soft, inconspicuous; perianth glabrous. ............... sects ataehedicds lating nou ane, behneutelama ukr eee ane doeap aie richocarpa. B. See usually erect; leaves oblong-elliptic, widest in eu Panes aes slightly rved; pollen n germinatio on aperture distinct and protru G. Shoots < . m in height; pseudostem and lower ae of leaf blade usually not glauc ; ee nie leaves held stiffly erect; cincinnal bracts red and yellow, margins involute at base; staminode apex apiculate; pollen with distal hemisphere lacking constrictions. .................. 6. H. marginata. H. Habitat terrestrial; leaves not held stiffly erect; cincinnal bracts entirely red, margins straight to revolute at base; staminode apex trilobed; pollen wi distal hemisphere having constrictions. I. Rachis glabrous, extremely a making inflorescence cae very contorted; perianth length 5-6 cm. ................. 7. H. secunda. . Rachis puberulous to tomentose, ere twisted so that ail cincin- se bracts oriented on one side of inflorescence; inflorescence usually highly contorted; perianth length 4-5 cm. ........ 8. H. nutans. G. isek. > 2 m in height; pseudostem and lower surface of leaf blade usually glaucous. J. Cincinnal bracts red and yellow; floral bracts aaa ovaries puber ulous to tomentose. ............0.. 00. c eee 9. H. playstachy J. Cincinnal bracts entirely red; floral bracts glabrous; ovaries glabrou sei ode Shae etree hae aaah aien Wee Waa at meee ees 10. H ae A. Inflorescences large (> 20 cincinnal bracts); cincinnal bract I/w < 1.4; anther rs con- nivent within perianth apex; pollen germination aperture distinct but sunken. K. Peduncle, rachis, and cincinnal bracts essentially glabrous; flowers parabolic or only slightly sigmoid; perianth essentially glabrous, sometimes slightly puberu- S) — cc s) S iS 3 =| cad io” 5 pee) ie) as 5 pe a: oo =| -O p pa Ae a = Vv a hod uo) oO t. janth y . Habitat terrestrial; lamina base truncate to ae aotuie long, inflo- 1984] KRESS, HELICONIA 463 rescence thus distinctly pendent; cincinnal bract 1/w 1.1-1.3; flowers par- abolic. N. Rachis slightly flexuose; cincinnal bracts distichous to spirally ar- ranged, alternate bracts distant and not touching. ................ Det P Non i Daa at ay cee ate Natg enon ee tare 11. H. curtispatha. N. Rachis very flexuose; cincinnal bracts strictly ao alternate bracts often touching or overlapping. .............. 12. H. stilesii. M. Habitat aquatic; lamina base ihe peduncle very short, obscure, inflorescence thus sessile and nodding to pendent; cincinnal bract 1/w < ; flowers slightly sigmoid. .......................0.. 13. H. sessilis. L. Cincinnal bracts imbricate, 1/w < 0.9; perianth red to pink. 14. H. mariae. Peduncle, rachis, and/or cincinnal bracts usually densely velutinous to woolly, in some cases all or some parts of sigmoid; perianth apex densely velutinous; pollen with ‘distal si tah truncate. O. Peduncle, rachis, and cincinnal bracts essentially glabrou P. Peduncle and rachis red or yellow, cincinnal bracts re red or red and yellow; floral bracts and perianth with golden hairs. ................. ns Sula Hoshasen ty ee wath ate cee dee asoee ae 15. H. pogonantha. P. Peduncle, rachis, and cincinnal bracts rose-red; floral bracts and perianth with bright yellow hairs. .....................05. 16. H. ramonensis. O. Peduncle, Been and/or cincinnal bracts densely velutinous, to woolly. Q. Cincinnal bracts bichromatic, red and yellow, velutinous. ............ dh oi ASS a eee a aia ee Be wn ee 15, i. pogonantha. Q. Saar en bracts monochromatic, not red and yellow, velutinous to woolly. orescence orange to rose-red, with orange hairs; perianth with ce or buff to rusty orange hairs S. Inflorescence pink to aie with rusty orange hairs; perianth with rusty orange hairs. .................. 1 . ramonensis. S. Inflorescence orange- oe with orange (fresh) 7 buff (dried) hairs. i eseatttacie aie ae one ae rea ace vas aerate ane sacs eet ocak? H. danielsiana. R. Inflorescence burgundy with golden to burgundy ae or yellow with yellow hairs; perianth with golden or yellow hairs. T. Inflorescence deep red to one with golden to burgundy hairs; n Iden hairs. ................. magnifica. T. Inflorescence bright ritee green with yellow hairs; ete with bright yellow hairs. .................... 19. H. xanthovillosa. 1. Heliconia necrobracteata Kress, J. Arnold Arbor. 62: 248. 1981. Type: anama, Coclé, 6 mi beyond El Valle de Antén, behind Cerro Gaital, elev. 900 m, Kress, Clarkson, & McDade 77-842 (holotype, DUKE!; isotypes, K!, Mo!). FiGurRE 8; PLate I, A, B. Herb with Musa-like habit, 4—4.5 m tall. Leafy shoots in groups of 3 to 12; pseudostem dull gray-green with brown spots, glabrous, 1-2 m tall, 4-6.5 cm in diameter; leaves 3 to 5 per shoot, all tending to lie in horizontal plane; longest petiole olive green, glabrous, 65-100 cm long, 1.5—2 cm in diameter; blade oblanceolate, widest toward acute to obtuse apex, the base unequal, attenuate along petiole, the upper surface dark green, glabrous, with midrib light green and glabrous, the lower surface gray-green, glabrous, with midrib yellow-green and glabrous, the longest blade 1.9-2.4 m by 25-35 cm. Inflo- rescence pendent, 40-55 cm long; peduncle red, glabrous to puberulous, 25- 35 cm long, 1.3—1.8 cm in diameter; rachis flexuose, dark red, puberulous, 1- 464 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 1.4 cm in diameter at base; cincinnal bracts spirally arranged, 10 to 20 per inflorescence, oriented 135-140° to axis of inflorescence, becoming more re- flexed with age, the basal bract usually fertile, the middle bract with apex acuminate, margins straight, inner surface yellow (sometimes with red margins) and glabrous, outer surface red and glabrous to puberulous, 12-17 cm long, 7-8 cm wide at base, I/w = 1.9, all bracts becoming necrotic soon after anthesis, deteriorating distally first and basipetally with age. Floral bracts more or less translucent at anthesis, quickly decomposing, white, puberulous to velutinous along abaxial margins, 3.5-4.5 cm long, 1.5—2 cm wide at base. Flowers 8 to 13 per cincinnus; pedicel pale yellow, tomentose, 1.4-2.2 cm long; perianth yellow to deep yellow, glabrous, puberulous along margins, 4.5-5 cm long, 6- 9 mm wide at base, at anthesis curved 50-80° and sigmoid; free sepal reflexed, fused sepals with apices slightly reflexed; staminode white, 1.5-2 cm by 4-5 mm, apex bilobed; stamens with anthers partially flared outside of apex of corolla tube; pollen trichocarpa type, medium size (54 x 74 wm), euoblate (P/E = 0.72), isopolar (DR/PR = 1.02), with the distal hemisphere planar, mi- nutely spinulose, the proximal hemisphere convex, minutely spinulose to gran- ulose, the germination aperture indistinct; ovary pale yellow, glabrous, 5-6 by 6-7 mm. Drupes glabrous. SPECIMENS EXAMINED. Panama. Coc e: base of three peaks beyond El Valle de Antén, elev. 800 m, Kress & Cooper-Smith 76-652 (DUKE); behind Cerro Gaital, elev. 2200 ft, Kress et al. 80-1158 (puKE), elev. 2400 ft, Kress & Hammel 83-1583 (DUKE); mtns. above EI Valle, in forest on Sr. Furlon’s Finca, Stimson 5037 (scz), 5038 (scz). PHENOLOGY. Flowering early in rainy season (April) through late August. HABITAT AND DISTRIBUTION. This species is found on steep, forested slopes at elevations of 650-1000 m in the region of El Valle de Anton in Panama (Map 1). As more collections are made in the provinces of Coclé and Veraguas, especially on the Atlantic slopes, the known distribution will probably be broad- ened RELATIONSHIPS. Heliconia necrobracteata, a member of the trichocarpa group, is distinctive in its glabrous ovaries, its stouter vegetative shoots and inflores- cences, and its red cincinnal bracts that decompose and turn black even while flowers are still being produced. 2. Heliconia maculata Kress, J. Arnold Arbor. 62: 244. 1981. Type: Panama, Col6n, along Rio Guanche, | km from road to Portobelo, elev. ca. 8 m, Kress & Knapp 80-1240 (holotype, DUKE!; isotypes, F!, GH!, K!, Mo!, PMA!). FiGure 9; PLATE I, C, D. Herb with Musa-like habit, 2—2.5 m tall. Leafy shoots congested, in groups of 3 to 25; pseudostem pale greenish gray with distinctive dark brown spots throughout, glabrous, 1.2-1.7 m tall, 2.2-2.5 cm in diameter; leaves 6 per shoot, all tending to lie in horizontal plane; petiole olive green, glabrous, 22- 34 cm long, 8-10 mm in diameter; blade oblanceolate, widest toward acute apex, the base unequal, attenuate, extending along petiole, the upper surface green, glabrous, with midrib light green and glabrous, the lower surface gray- green, glabrous, with midrib yellow and glabrous to slightly scurfy, the longest 1984] KRESS, HELICONIA 465 Figure 8. Heliconia necrobracteata. A, inflorescence. B, C, cincinnal bracts: B, entire a, flower protruding at anthesis); C, cut-away, floral bracts removed showing flower G, abaxial view; H, position relative to style, lateral view. I, style and stigma. blade 1.1-1.3 m by 24-26 cm. Inflorescence pendent, to 51 cm long; peduncle green and dark maroon, glabrous, 7-32 cm long, | cm in diameter; rachis flexuose, dull red, glabrous to slightly puberulous, 7-8 mm in diameter at base; cincinnal bracts spirally arranged, 7 or 8 per inflorescence, oriented 90° to axis 466 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 T T T T ae 86 82 78 1 i i l AP Costa Rica and Panama, showing allopatric distribution of Heliconia nec- robracteata (triangles), H. maculata (circles), and H. talamancana (squares). of inflorescence, the basal bract fertile or sterile, the middle bract with apex long-acuminate, margins straight becoming somewhat flared at base, inner surface pale orange and glabrous, outer surface red basally becoming green toward apex and glabrous, 14-17 cm long, 4—4.5 cm wide at base, l/w = 3.7. Floral bracts translucent at anthesis, quickly decomposing, cream, glabrous to puberulous abaxially, 4-5 cm long, 1.1-1.4 cm wide at base. Flowers 15 to 20 per cincinnus; pedicel pale yellow, pubescent to hirsute, 1—1.5 cm long; perianth yellow basally, green at apex, glabrous to slightly puberulous, 5 cm long, | cm wide at base, at anthesis curved 30-50° and slightly sigmoid; free sepal slightly reflexed, fused sepals with apices not reflexed; staminode white, 7-10 by 3 mm, apex apiculate; stamens with anthers flared outside perianth apex; pollen trichocarpa type, medium size (49 x 67 um), euoblate (P/E = 0.73), isopolar (DR/PR = 0.97), the distal hemisphere subconvex, verrucose, the proximal hemisphere convex, granulose, the germination aperture indistinct; ovary white, pubescent, 4-5 by 6 mm. Drupes glabrous to puberulous, 8 by 9 mm; pyrenes 6 by 5 mm. SPECIMENS EXAMINED. Panama. CoLon: Rio Guanche, forest, elev. ca. 50 ft, Maas et al. 1583 (F, Mo); in forest 3-7 km from bridge, elev. 300-700 ft, Hammel et al. 4899 (mo); Knapp 1020 (mo), 1422 (mo); below Cerro Bruja along Rio Escondaloso, elev. 100-200 m, Kress & Knapp 82-1406 (DUKE). PHENOLOGY. Flowering late July through early October. HABITAT AND DISTRIBUTION. This species occurs at lower elevations along stream banks and in adjacent understory of primary forest. It will invade more open areas (often created by man), where it generally produces more shoots per clump. So far, Heliconia maculata has been collected at only two localities in Col6n Province, Panama (Map 1); as more collections are made, it will probably be found in other forested areas in the Atlantic coastal forests of Panama. 1984] KRESS, HELICONIA 467 D, E, floral bracts: D, abaxial view; E, lateral view. F, perianth, outer surface showing 2 sepals fused to partially spread-open corolla tube. G, H, staminode: G, abaxial view; H, position relative to style, lateral view. I, style and stigma. J, pseudostem. K, vegetative habit. 468 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 RELATIONSHIPS. Several pollen characters, as well as the distinctive pubescent ovaries and the obliquely or horizontally oriented leaf blades that are widest near the apex, ally this species with other members of the trichocarpa group. However, it differs from the other species in its dark brown-spotted pseudostem (also present in Heliconia necrobracteata), its red and green cincinnal bracts, and its yellow and green, glabrous perianths. 3. Heliconia talamancana Daniels & Stiles, Brenesia 15(Supl.): 42. 1979. Type: Costa Rica, [Lim6n,] on ridge W of BriBri, elev. 200 m, Daniels, Stiles, & Kress 115 (holotype, F!; isotypes, CR, US). PLATE I, E, F. Herb with Musa-like habit, 4-4.5 m tall. Leafy shoots in groups of 5 to 15; pseudostem green, glabrous, 2.2—2.6 m tall, 3.5—4 cm in diameter; leaves 5 per shoot, all tending to lie in horizontal plane; petiole olive green, glabrous, 85- 100 cm long, 1.5-1.6 cm in diameter; blade oblanceolate, widest toward obtuse apex, the base unequal, attenuate along petiole, the upper surface green, gla- brous, with midrib green and glabrous, the lower surface gray-green, glabrous, with midrib green-yellow and glabrous, the longest blade 2.1-2.5 m by 32-35 cm. Inflorescence pendent, to 65 cm long; peduncle green, glabrous, 6-27 cm long, 1.3-1.5 cm in diameter; rachis flexuose, green and red, glabrous, 1.2-1.4 cm in diameter at base; cincinnal bracts spirally arranged, 8 to 13 per inflo- rescence, oriented 90° to axis of inflorescence, the basal bract usually fertile, the middle bract with apex long-acuminate, margins straight, inner surface white to green and glabrous, outer surface green becoming red at rachis and glabrous, 17-20 cm long, 6-7 cm wide at base, I/w = 2.9. Floral bracts opaque, decomposing after anthesis, white, glabrous to puberulous abaxially, 5-5.3 cm long, 1.5—2 cm wide at base. Flowers 15 to 20 per cincinnus; pedicel white, tomentose, |.3-1.6 cm long; perianth white, green apically, puberulous, 5—5.2 cm long, | cm wide at base, at anthesis curved 30-40° and slightly sigmoid; free sepal reflexed, fused sepals with apices not reflexed; staminode white, spatulate, 1.1 cm by 3 mm, apex apiculate; stamens with anthers partially flared outside corolla tube; pollen trichocarpa type, medium size (53 x 72 um), euob- late (P/E = 0.74), isopolar (DR/PR = 0.95), with the distal hemisphere sub- convex to planar, verrucose to granulose, the proximal hemisphere convex, verrucose to granulose, the germination aperture ee aad white, vil- lous, 6-8 by 7-8 mm. Drupes puberulous, 1-1.2 by 1.2-1.5 ¢ SPECIMENS EXAMINED. Costa Rica. Limon: 1-3 km N of BriBri, Rio Sixaola Drainage, elev. 20-200 m, Burger & Antonio 11000 (F); in hills above BriBri, elev. ca. 250 m, Kress et al. 76- 625 (DUKE), elev. ca. 800 ft, Kress et al. 77-791 (DUKE). Panam a, VE- RAGUAS: along road on Pacific slope, 1-3 km above Escuela Agricola Alto ae elev. 700-800 m, Croat 25997 (mo) PHENOLOGY. Flowering June to October. HABITAT AND DISTRIBUTION. This species is distributed on the Caribbean slopes from the Rio Sixaola in Limon, Costa Rica, to Veraguas, Panama (Map 1). It is most often found on steep, wooded slopes in primary and old secondary growth at elevations up to 700 m 1984] KRESS, HELICONIA 469 RELATIONSHIPS. Heliconia talamancana is closely related to H. maculata and other species in the trichocarpa group. It differs from H. maculata in its greater overall stature (> 3 m), its essentially green cincinnal bracts, and its green, pubescent perianth. 4, Heliconia colgantea R. R. Smith ex Daniels & Stiles, Brenesia 15(Supl.): 18. 1979. Type: Costa Rica, [Puntarenas,] 10 km ENE of Palmar Norte, Burger & Matta 4662 (holotype, F!; isotypes, cR!, GH!, Ny!, Us!). PLATE II, A, C. Herb with Musa-like habit, 2 m tall. Leafy shoots solitary or in pairs; pseu- dostem yellow-green, glabrous, 90-125 cm tall, 2.2-2.7 cm in diameter; leaves 6 per shoot, all fanning out from pseudostem and tending to lie in horizontal lane; petiole green to dark green, glabrous, 34-45 cm long, 9-11 mm in diameter; blade oblanceolate, widest toward obtuse apex, the base unequal, obtuse to attenuate along petiole, the upper surface dark green, glabrous, with midrib green and glabrous, the longest blade 1.1-1.3 m by 20-25 cm. Inflo- rescence pendent, to 45 cm long; peduncle red with green striations, puberulous, 14-17 cm long, |-1.2 cm in diameter; rachis flexuose, pink-red, puberulous to tomentose, 9-10 mm in diameter at base; cincinnal bracts spirally arranged, 8 to 10 per inflorescence, oriented 120-—165° to axis of inflorescence, becoming more reflexed with age, the basal bract usually fertile, the middle bract with apex acuminate, margins straight becoming somewhat revolute near rachis, inner surface yellow and pink and glabrous, outer surface pink becoming green toward apex and puberulous, 19-22 cm long, 7-8 cm wide at base, I/w = 2.7. Floral bracts translucent, persistent, conspicuous, pale yellow, stiff, puberulous abaxially, 3.5-4.2 cm long, 8-10 mm wide at base. Flowers 15 to 20 per cincinnus; pedicel pale yellow, puberulous to tomentose, !-1.2 cm long; peri- anth yellow, densely puberulous, 5.1-5.5 cm long, 8-9 mm wide at base, at anthesis curved 50—70° and sigmoid; free sepal reflexed, fused sepals with apices sometimes slightly reflexed; staminode white, 1-1.1 cm by 2-3 mm, apex bidentate; stamens with anthers flared outside perianth apex; pollen ¢richocarpa type, medium size (55 x 79 wm), euoblate (P/E = 0.69), isopolar (DR/PR = 0.88), with the distal hemisphere planar, minutely spinulose, the proximal hemisphere convex, minutely spinulose to granulose, the germination aperture indistinct; ovary pale yellow, puberulous to tomentose, 4-6 by 6-7 mm. Drupes sparsely tomentose. SPECIMENS EXAMINED. Costa Rica. PUNTARENAS: forested hills near Palmar Norte, elev. 300 m, Allen 6551] (F, PMA); 10 km N of Palmar Norte along Pan Am Hwy., elev. ca. 100 m, Kress et al. 76-595 (DUKE), 77-878 (DUKE); cult. at Las Cruces Trop. Bot. Garden, San Vito de Java, elev. 3900 ft, Kress 78-1027 (DUKE). Panama. CHIRIQUi: Burica Pen- insula, 10-11 mi W of Puerto Armuelles in vic. of San Bartolo Limite, elev. 300-500 Pucuro base camp, elev. 650 m, Gentry & Mori 13877 (mo); E slope of Cerro Sapo, elev. 2500 ft, Hammel 1275 (mo); Rio Pirre, near Dos Bocas, Foster & Augsburger 2844 (F); Cerro Pirre, Bristan 628 (Ny), 629 (Ny); 4.5 km S of El Real, moist forest at base of foothills of N slope of Cerro Pirre, Mori & Kallunki 5413 (GB, Mo); headwaters of Rio 470 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Tuquesa, vic. of gold-mining camp of Tyler Kittredge, Croat 27198 (mo), 10 km NE of Jaqué, Rio Pavarando, elev. 1400 ft, D’Arcy & Sytsma 14522 (Mo), Sytsma & D'Arcy 3375 (Mo); Rio Jaqué valley, elev. 300-500 m, Knapp & Mallet 3085 (Mo). PANAMA: S of Ipeti, Serrania de Majé, elev. 450-600 m, Knapp et al. 4553 (mo). PHENOLOGY. Flowering mainly January to September. HABITAT AND DISTRIBUTION: Heliconia colgantea has been found in several localities in Costa Rica and Panama (Map 2). It is restricted mainly to moist or wet forested slopes and ravines between 100 and 800 m. RELATIONSHIPS. This species is a member of the sis group. Slightly pubescent, bright pink cincinnal bracts with green apices, and stiff, conspicuous floral bracts distinguish Heliconia colgantea ‘from its closest relative, H. trichocarpa. 5. Heliconia trichocarpa Daniels & Stiles, Brenesia 15(Supl.): 44. 1979. Type: Costa Rica, Alajuela, La Balsa area N of San Ramon, elev. 800 m, Daniels, Kress, & Hutchison 112 (holotype, F!; isotypes, CR, US). PiateE II, B, D. H. trichocarpa var. caducispatha Daniels & Stiles, Brenesia 15(Supl.): 44. 1979. Type: Costa Rica, Puntarenas, 42 km S of Palmar Norte along Interamerican Hwy., elev 100 m, Daniels & Stiles 129 (holotype, F!; isotypes, CR, US) Herb with Musa-like habit, 2-4 m tall. Leafy shoots in groups of 4 to 20; pseudostem green (sometimes with red), glabrous, 80-190 cm tall, 2-4 cm in diameter; leaves 4 to 6 per shoot, all tending to lie in horizontal plane; petiole green to olive green, glabrous, 35-90 cm long, 7-13 mm in diameter; blade oblanceolate, widest toward acute to obtuse apex, the base unequal, obtuse to attenuate along petiole, the upper surface green to dark green, glabrous, with midrib light green to olive green and glabrous, the lower surface gray-green, glabrous, with midrib yellow-green and scurfy, the longest blade 85-175 by 17-30 cm. Inflorescence pendent, to 60 cm long; peduncle red to rose-red, puberulous, 10-50 cm long, 1-1.8 cm in diameter; rachis flexuose, red, pu- berulous, 6-11 mm in diameter at base; cincinnal bracts spirally arranged, 6 to 10 per inflorescence, oriented 110-150° to axis of inflorescence, the basal bract usually fertile, the middle bract with apex long-acuminate, margins straight, inner surface white to yellow to pink and glabrous, outer surface red to rose- red and glabrous to puberulous, 12-28 cm long, 6-8 cm wide at base, 1/w = 2.9. Floral bracts translucent, persistent, inconspicuous, white to yellow, soft, puberulous abaxially, 4.5-6 cm long, 1.5-2.5 cm wide at base. Flowers 10 to 20 per cincinnus; pedicel pale yellow, puberulous to tomentose, 7-13 mm long; perianth yellow, glabrous to puberulous, 4.6-5.5 cm long, 1-1.2 cm wide at base, at anthesis curved 50-70° and sigmoid; free sepal reflexed, fused sepals with apices not reflexed; staminode white, 1-1.3 cm by 2-3 mm, apex bidentate; stamens with anthers flared outside perianth apex; pollen ftrichocarpa type, medium size (53 x 78 um), euoblate (P/E = 0.68), isopolar (DR/PR = 1.07), with the distal hemisphere planar, minutely spinulose to verrucose, the prox- imal hemisphere convex, minutely spinulose to granulose, the germination 1984] KRESS, HELICONIA 471 T Ie T T T @ e e® «e 5 a 1044 eo? e a v v v 7 ae ‘ % e at v ¥, & & * v fil Vv v Vv v ye i) 8 6 j 82 f it p 2. Costa Rica and Panama, showing distribution of Heliconia colgantea (tri- es and H. trichocarpa (circles). aperture indistinct; ovary white to pale yellow, puberulous to tomentose, 5-8 by 6-9 mm. Drupes tomentose. SPECIMENS EXAMINED. Costa Rica. ALAJUELA: entre Cataratas y La Balsa de San Ramon, elev. 700-800 m, Brenes 4429 (cr, F), elev. ca. 1000 m, Kennedy 1657 (mo), elev. ca. O m, Kress et al. 76-605 (DUKE), elev. 2700 ft, Kress et al. 77-767 (DUKE), 77-807 (DUKE), elev. 900-1000 m, Stevens 13791 (mo); Finca Los Ensayos ca. 11 mi NW of Zarcero, elev. 850 m, Croat 43597 (mo); Buena Vista de San Carlos, Quebrada Lajas, Finca Los Ensayos, elev. 850 m, Jiménez 2323 (cR, F, NY); between San Lorenzo and Los Angeles de San Ram6n, above Rio San Lorenzo, elev. 620 m, Burger & Antonio 11190 (F); Colonia Virgen del Socorro, barranca of Rio Sarapiqui, elev. 700-800 m, Stevens 13565 (Mo). CarTAGo: foréts de Tuis, elev. 650 m, 76nduz 11383 (cr, us); woods above Rio Pejibaye, 2 km SW of Taus, elev. 740 m, Lent 2986 (F); near Moravia, elev. 1150-1200 m, Maas 1075 (uv). CARTAGO/LIMON: between Turrialba and Siquirres, elev. 2000-2500 ft, Kress et al. 77-787 (DUKE). GUANACASTE: below Volcan Tenorio along road to Upala, elev. 700 ft, Kress et al. 80-1217 (DUKE). HEREDIA: Finca La Selva near Puerto Viejo de ao me ca. 150 m, Kress 79-1109 (DUKE). LIMOn: along Hwy. 32 from Turrialba to Limén, 11 mi S of Siquirres, elev. 650 m, Croat 43350 (mo). Kress 76-572 (DUKE), 78-975 (one), SAN José: streamsides along Rio Claro Valley (Rio La Hondura drainage), below La Palma, NE of San Jer6nimo, elev. 1000-1200 m, Burger & Burger 7657B (F). Panama, CuHirRiQui: Fortuna, elev. 1000-1200 m, Correa et al. 2668 (mo); upriver from main camp, Fortuna Dam Site, 1200-1400 m, Folsom et al. 5509 (mo); Kress & Hammel 83-1605 (DUKE). CocLé: El Copé, beyond sawmill, elev. 2800 ft, Kress et al. 80-1171 (puKE), Kress & Hammel 82-1323 (DUKE); El Potroso, elev. 800- 0 m, Andersson & Sytsma 1276 (s), D’Arcy 11273 (Mo). VERAGUAS: valley of Rio Dos Bocas on road between Alto Piedra (above Santa Fé) and Calovébora, elev. 350- 400 m, Croat 27429 (mo), elev. 500 m, Andersson & Sytsma 1286 (GB, 8); beyond Santa Fé above Escuela Agricola, elev. 2600 ft, Kress & Cooper-Smith 76-657 (DUKE), Kress et al. 77-832 (DUKE); Atlantic slope beyond Santa Fé on road to Calovébora, elev. 1200 472 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 ft, Kress et al. 80-1196 (puKE); Rio Calabacito, Aguabal, elev. 400 m, Maas & Dressler 1624 (MO, U). PHENOLOGY. Flowering mainly during late rainy season (July to November), peaking September and October. HABITAT AND DISTRIBUTION. Heliconia trichocarpa is distributed throughout Costa Rica and Panama, primarily on the Atlantic slopes at elevations between 100 and 1000 m (Map 2). This species is found in a wide range of habitats, from stream margins in primary forest understory to roadside embankments in early secondary growth. VARIATION. Daniels and Stiles (1979) recognized two varieties of Heliconia trichocarpa: var. trichocarpa and var. caducispatha. They listed several char- acters by which the two taxa differ, the most important one being the rapid postanthesis disintegration of the cincinnal bracts in the latter variety. Although there are some morphological differences, the extent of variation between va- rieties is not greater than that between populations of var. trichocarpa on the Atlantic slopes. In addition, intermediate forms between varieties are known (e.g., Kress et al. 76-592, puke). For these reasons var. caducispatha is not accepted here. RELATIONSHIPS. Heliconia trichocarpa is allied to other species of the tricho- carpa group with horizontally oriented oblanceolate leaf blades that are widest toward the apex, smaller pendent inflorescences with long-acuminate cincinnal bracts (I/w > 2.5), pubescent ovaries, and pollen with indistinct germination apertures. Its closest relative is H. colgantea, from which it differs in its gla- brous, red cincinnal bracts, its inconspicuous, decomposing floral bracts, and its glabrous perianth. 6. Heliconia marginata (Griggs) Pittier, Man. Pl. Usual. Venez. 299. 1926. PLateE III, A, B. Bihai marginata Griggs, Bull. Torrey Bot. Club 42: 323. 1915. Type: Panama, [Darién,] Marraganti and vic., Williams 696 (holotype, Ny!; isotype, us!). Herb with Musa-like habit, 2-3 m tall. Leafy shoots in groups of 20 to over 50; pseudostem gray-green, glabrous, 45-120 cm tall, 4.3-7 cm in diameter; leaves 4 or 5 per shoot, held stiffly erect; petiole green, glabrous, 45-90 cm long, 8-12 mm in diameter; blade with the base unequal, obtuse to attenuate, the apex acuminate, the upper surface green to bright green, glabrous, with midrib green and glabrous, the lower surface green, glabrous to slightly glaucous, with midrib green and glabrous, the longest blade 60-120 by 17-25 cm. Inflo- rescence pendent, to 40 cm long; peduncle red and green, scurfy, 35-60 cm long, 9-10 mm in diameter; rachis flexuose, red, puberulous to scurfy, 6-12 mm in diameter at base; cincinnal bracts distichous to spirally arranged, 9 to 14 per inflorescence, oriented 90-120° to axis of inflorescence, the basal bract often sterile, the middle bract with apex acute to acuminate, margins straight becoming involute at base, inner surface yellow and glabrous, outer surface red becoming yellow along margins and puberulous, 7—9.5 cm long, 4.5-5.5 cm 1984] KRESS, HELICONIA 473 wide at base, I/w = 1.7. Floral bracts opaque, persistent, yellow and pink, puberulous to villous abaxially, 5-6 cm long, 1.9-2 cm wide at base. Flowers 7 to 15 per cincinnus; pedicel pale yellow with pink striations, puberulous, 1- 1.3 cm long; perianth yellow, glabrous, puberulous along margins, 4.9-5.4 cm long, 8-10 mm wide at base, at anthesis curved 35-45°; free sepal strongly reflexed, fused sepals with apices not reflexed; staminode white, 6-8 by 3-4 mm, apex apiculate, base constricted; stamens with anthers connivent inside apex of corolla tube or slightly flared outside perianth apex; pollen nutans type, to subconvex, psilate to granulose, the germination aperture distinctly pro- truding; ovary pale yellow, glabrous, 9-10 by 5-6 mm. Drupes glabrous, |.6- 1.9 by 1-1.3 cm SPECIMENS EXAMINED. Costa Rica. PUNTARENAS: 8 mi from Quepos on road to Parrita, ca. sea level, Kress et al. 77-804 (DUKE); Finca 44 near Rio Colorado beyond Villa Neily, elev. ca. 50 ft, Kress & Cooper-Smith 76-661 (DUKE), Kress & Clarkson 79-1095 (DUKE); near Coto 47 in herbaceous swamp, sea level, Maas & McAlpin 1465 (cr, vu). Panama. Darién: Alconorque swamp on Rio Tuira ca. 3 mi NW of El Real, Duke 4817 (Gu, us); .5 mi E of airstrip at El Real, Lazor & Correa 3398 (scz); Cerro Tacarcuna-Serrania, near Yaviza, elev. 50 m, Gentry & Mori 13506 (Mo). PHENOLOGY. Flowering and fruiting nearly all year, peaking July and August. HABITAT AND DISTRIBUTION. This species is found on the Pacific Coastal Plain of Costa Rica and Panama, extending north to the Quepos area and south into Darién and South America, where it is more common (see Map 3). It almost always occurs in dense stands in open swamps or standing water. Heliconia marginata is one of the few truly aquatic species of the genus. RELATIONSHIPS. Among Central American species with pendent inflorescences, Heliconia marginata is most closely allied to members of the nutans group. However, differences in cincinnal bract morphology, floral structure, and pollen characteristics suggest that this species probably has closer relatives among South American taxa. The small inflorescences with red and yellow cincinnal bracts, the aquatic habitat, and the stiff, erect leaf blades readily distinguish H. marginata from other Central American heliconias. 7. Heliconia secunda R. R. Smith, Phytologia 30: 214. 1975. Type: Costa Rica, Heredia, along steep slope near road and waterfall, ca. 7 km N of Vara Blanca de Sarapiqui, elev. 5000 ft, R. R. Smith 2206 (holotype, FLAS; isotypes, A!, GH, MO, US Herb with Musa-like habit, 2-3 m tall. Leafy shoots solitary or in groups of 2 to 50; pseudostem glabrous, 0.6—2 m tall, 2.5—4 cm in diameter; leaves 2 to 5 per shoot; petiole green, glabrous, 40-120 cm long, 0.8-1.3 cm in diameter; blade with the base unequal, obtuse, the apex acute, the upper surface green, glabrous, with midrib green and glabrous, the lower surface light green to maroon, glabrous, with midrib green to yellow-green and maroon and glabrous to scurfy, the longest blade 80-120 by 20-40 cm. Inflorescence nodding to 474 JOURNAL OF THE ARNOLD ARBORETUM (VOL. 0) T a T T T T Ye i ra a : fe 144 ; e® Y L ~ 124 v vy oe e + bg 10-4 N eo & 7 86 B2¢e 78 t L 1 | 1 Map Nicaragua, Costa Rica, and Panama, showing distribution of Heliconia mar- ginata (squares), H. secunda vars. secunda (circles) and viridiflora (triangles). pendent (sometimes contorted), to 50 cm long; peduncle red to yellow-red and green, glabrous to scurfy, 3.5—25 cm long, 1-1.3 cm in diameter; rachis flexuose and twisted or contorted, red, glabrous to scurfy, 9-1 1 mm in diameter at base; cincinnal bracts distichous or spirally arranged, 8 to 12 per inflorescence, ori- ented 90-1 20° to axis of inflorescence, the basal bract usually fertile, the middle bract with apex acuminate, margins straight to revolute, inner surface red to orange and glabrous, outer surface red and glabrous to slightly scurfy, 10-14 cm long, 5.3-8 cm wide at base, I/w = 1.8. Floral bracts translucent, slowly decomposing after anthesis, pale yellow, glabrous to puberulous along midrib abaxially, 4-6 cm long, 1.5—2.3 cm wide at base. Flowers 10 to 25 per cincinnus; pedicel white to green to pale yellow, glabrous, 5—10 mm long; perianth yellow to green, glabrous except puberulous along sepal margins, 5-6 cm long, 7-10 mm wide at base, at anthesis curved 20—50°; free sepal reflexed, fused sepals with apices not reflexed; staminode white, apex 7-10 by 3-4 mm, tridentate with center tooth longer than blunt lateral lobes; stamens with anthers flared outside perianth apex; pollen nutans type, small (43-44 x 58-62 um), euoblate (P/E = 0.70-0.74), subisopolar (DR/PR = 2.24-2.55) with the distal hemi- sphere constricted, convex, spinulose, the proximal hemisphere planar to sub- convex, psilate to verrucose, the germination aperture distinctly protruding; ovary light green to yellow, glabrous, 6-9 by 5-6 mm. Drupes glabrous. DistRIBUTION. Heliconia secunda is found in middle-elevation forests of Costa Rica and Nicaragua (Map 1984] KRESS, HELICONIA 475 VARIATION. This species consists of two varieties that differ primarily in flower color. RELATIONSHIPS. Heliconia secunda, a member of the nutans group, is closely related to H. nutans (with pendent inflorescences) and H. tortuosa (with erect inflorescences). It is distinguished from other members of the group by its extremely twisted rachis, which gives the nodding to pendent inflorescences a very contorted appearance. Key to the Varieties of Heliconia secunda Leaf blade with lower surface green; perianth yellow; ovary pale green. .............. Bees Saket Reh ay SR te he Es dations to carn tad atta eae 7a. var. los Leaf blade with lower surface usually maroon; perianth green; ovary yellow to green. ee Gs ah sesh aha swe ashe d eeeshaeeg agnan sees oncieacdvice. ay taste ees 7b. var. viridiflora 7a. Heliconia secunda var. secunda PLATE III, C, D. Leafy shoots in groups of 10 to 50; pseudostem |.4—2 m tall, 2.5-4 cm in diameter; leaves 2 or 3 per shoot; petiole 50-100 cm long, 1—1.3 cm in diameter; longest blade 80-120 by 30-40 cm. Inflorescence nodding to pendent, to 50 cm long; peduncle red to yellow-red and green, scurfy, 8-25 cm long, 1-1.3 cm in diameter; rachis scurfy, 9-11 mm in diameter at base; cincinnal bracts spirally arranged, 9 to 12 per inflorescence, oriented 90—120° to axis of inflo- rescence, the middle bract with margins revolute, inner surface red becoming yellow at midrib, 10-12.5 cm long, 5.3-6 cm wide at base. Floral bracts 4-6 cm long, 1.5-2.3 cm wide at base. Flowers 15 to 25 per cincinnus; pedicel green, 5-10 mm long; perianth yellow, 5.5—6 cm long, 8-10 mm wide at base, at anthesis curved 20-30°; staminode 9-10 by 3 mm; ovary light green, 6-9 by 5-6 mm. Drupes 1.4 by | cm SPECIMENS EXAMINED. Nicaragua. JINOTEGA/MATAGALPA: in forest beside hwy. between Jinotega and Matagalpa, elev. 4000-4500 ft, Bunting & Licht 981 (F, Ny, US). JINOTEGA: region of Las Mercedes, sierra E of Jinotega, chiefly in dense wet mixed virgin forest (cloud forest), elev. 1200-1500 m, Standley 10724 (F); Hwy. 3, 1.9 km NW of Aranjuez road entrance, elev. 1460-1480 m, Stevens 9202 (mo). Costa Rica. HEREDIA: Catarata E] Angel, Rio La Paz Grande entre Cariblanco y Vara Blanca, elev. 1350 m, Jiménez 2048 (F); at waterfall 3 mi from Cinchona, elev. 4100 ft, Kress & Cooper-Smith 76-529 (DUKE), elev. 4400 ft, Kress et al. 77-776 (DUKE); vic. of Vara Blanca, elev. 1750-1800 m, Luteyn 3021] (DUKE); Cinchona, along road to alee near waterfalls, Sheffy 68 (cR). SAN Jose: Rio Claro Valley, below La Palma, NE of Sa (Rio La Hondura drainage), elev. 1000-1200 m, Burger & Burger 76574 (F), cp 1500 m, Burger & Stolze 5303 (F), Burger & Visconti 10230 (F); Alto La Palma, elev. 4700 ft, Kress et al. 77-773 (DUKE); along road between La Palma and La Hondura, Sheffy 74 (cr); between Cascajal and San Isidro de Coronado, elev. 1600-1700 m, Taylor 935 (DUKE). PHENOLOGY. Flowering primarily March to August. HABITAT AND DISTRIBUTION. Heliconia secunda var. secunda has been collected only around Jinotega in Nicaragua and in several localities in the Cordillera Central of Costa Rica (Map 3). This variety inhabits old secondary growth in moist cloud forest at elevations of 1200-1500 m. 476 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 7b. Heliconia secunda var. viridiflora Daniels & Stiles, Brenesia 15(Supl.): 42. 1979. Type: Costa Rica, Alajuela, ca. 2 km N of Bijagua, elev. 350 m, Stiles s.n., 10 Sept. 1977 (holotype, uss; isotypes, F!, US). Heliconia viridiflora (Daniels & Stiles) Stiles, Brenesia 18: 152. 1980. Leafy shoots solitary or in groups of 2 to 4; pseudostem green, glabrous, 60— 150 cm tall, 3-4 cm in diameter; leaves 5 per shoot; petiole 40-120 cm long, 8-10 mm in diameter; blade with base slightly attenuate to obtuse, lower surface green (becoming maroon at higher elevations), longest blade 90-120 by 20-30 cm. Inflorescence nodding or contorted, to 36 cm long; peduncle red, glabrous, 3.5-5 cm long, 1-1.2 cm in diameter; rachis flexuose and contorted, red, gla- brous, | cm in diameter at base; cincinnal bracts distichous and often all arranged on same side of rachis, 8 to 10 per inflorescence, oriented 90° to axis of inflorescence, the middle bract with margins straight (sometimes overlapping near rachis), inner surface orange becoming red along margins, outer surface glabrous, 11-14 cm long, 7-8 cm wide at base. Floral bracts 4—4.5 cm long, 1.5-2 cm wide at base. Flowers 10 to 15 per cincinnus; pedicel pale yellow, 5- 8 mm long; perianth white basally, green distally, 5-5.5 cm long, 7-9 mm wide at base, at anthesis curved 35—50°; staminode 7-10 by 3-4 mm; ovary light green to yellow, 8 by 5-6 mm wide. Fruits not seen. SPECIMENS EXAMINED. Nicaragua. ZELAYA: ca. 6.3 km S of bridge at Colonia Yolaina and ca. 0.8 km S of ridge of Serrania de Yolaina on road to Colonia Manantiales (Colonia Somoza), elev. ca. 200-300 m, Stevens 488] (Mo). Costa Rica. ALAJUELA: 3 km NNE of Bijagua along new road to Upala, elev. 450 m, Burger & Baker 9815 (F), 9873 (F); wet forest 5 km S of Canalete near Rio Zapate, along new road to Upala, elev. 100-200 m, Burger & Baker 10009 (Ff). GUANACASTE: below Volcan Tenorio along road to Upala, elev. 800 ft, Kress et al. 80-1216 (DUKE), elev. ca. 700 ft, Kress et al. 80-1229 (DUKE). PUNTARENAS: San Vito de Java, cult. at Las Cruces Trop. Bot. Garden, elev. ca. 4000 ft, Kress 78-1033 (DUKE). PHENOLOGY. Flowering primarily August to November. HABITAT AND DISTRIBUTION. Heliconia secunda var. viridiflora 1s found in the Cordillera de Yolaina of Nicaragua and the Cordillera de Guanacaste of Costa Rica between 200 and 500 m (Map 3). This variety inhabits both primary and secondary forests. RELATIONSHIPS. Variety viridiflora differs from the typical variety of Heliconia secunda in its flowers with green perianths, and its leaf blades with the lower surfaces varying from green with maroon borders to totally maroon. Because of the nonoverlapping flowering times of the two varieties, Stiles (1980) rec- ognized this taxon as a species distinct from H. secunda. This phenological difference is not accepted here as sufficient evidence for elevating var. viridiflora to specific rank. 8. Heliconia nutans Woodson, Ann. Missouri Bot. Gard. 26: 276. 1934. TyPE: Panama, Chiriqui, Volcan de Chiriqui, vic. of Casita Alta, Woodson, Allen, & Seibert 968 (holotype, Mo!; isotypes, GH!, MICH, NY!, Us!). Ficure 10; PLaAte III, E, F. 1984] KRESS, HELICONIA 477 H, position relative to style, lateral view. I, style and stigma. J. inflorescence, schematic showing alternative orientation with all cincinnal ee aligned on | side of alorecene: axis. 478 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Heliconia allenii Standley & Williams, Ceiba 3: 189. 1953. Type: Panama, Chiriqui, Rio Chiriqui Viejo, vic. Nueva Suiza, elev. 5000 ft, Allen 6255 (holotype, us!). Heliconia villosa auct. non Klotzsch: Woodson & Schery, Ann. Missouri Bot. Gard. : 53. 1945, in part (Woodson and Schery included H. irrasa, H. tortuosa, and H. nutans in H. villosa). Herb with Musa-like habit, 1-2.5 m tall. Leafy shoots in groups of 5 to 30; pseudostem light green and maroon, glabrous, 50-160 cm tall, 1.5-3.8 cm in diameter; leaves 3 to 5 per shoot; petiole green and maroon, glabrous, 20-68 cm long, 5—1 1 mm in diameter; blade with the base unequal, obtuse to truncate, the apex acute, the upper surface dark green, glabrous, with midrib green and glabrous, the lower surface green to maroon, glabrous, with midrib light green to maroon and glabrous to scurfy, the longest blade 50-105 by 15-31 cm. Inflorescence nodding to pendent (sometimes contorted), to 55 cm long; pe- duncle green and red to solid red; puberulous to tomentose with golden hairs, 5-50 cm long, 5-10 mm in diameter; rachis flexuose (sometimes twisted), red to orange-red (sometimes yellow), puberulous to tomentose with golden hairs, 5-10 mm in diameter at base; cincinnal bracts spirally arranged to distichous (sometimes all bracts aligned on 1 side of inflorescence axis), 5 to 13 per inflorescence, oriented 90-135° to axis of inflorescence, the basal bract often sterile, the middle bract with apex acute to acuminate, margins straight to revolute distally, inner surface red to red-orange and yellow and glabrous to puberulous, outer surface red to red-orange and glabrous to puberulous, 7-11 cm long, 4-7.5 cm wide at base, I/w = 1.6. Floral bracts translucent, persistent and papery when dried or decomposing after anthesis, pale yellow, glabrous to puberulous along midrib abaxially, 3.5—4.5 cm long, 1.3—2 cm wide at base. Flowers 10 to 20 per cincinnus; pedicel white to yellow, glabrous, 4-10 mm long; perianth yellow, glabrous to slightly puberulous, 4-5 cm long, 6-9 mm wide at base, at anthesis curved 30-—50°: free sepal straight to reflexed, fused sepals with apices not reflexed; staminode white, apex 7-10 by 3-5 mm, trilobed with rounded lateral lobes; stamens with anthers flared outside perianth apex; pollen nutans type, small (46 x 54 um), suboblate (P/E = 0.87), subisopolar (DR/PR = 2.49) with the distal hemisphere constricted, convex, spinulose, the proximal hemisphere planar to subconvex, psilate to verrucose, the germination pore distinctly protruding; ovary light green, glabrous, 6-7 by 5-6 mm. Drupes glabrous, |-1.1 by 1-1.1 cm SPECIMENS EXAMINED. Costa Rica. PUNTARENAS: Las Alturas and vic., elev. 1600-1800 m, Maas & McAlpin 1486 (Ny, u); San Vito de Java, cult. at Las Cruces Trop. Bot. Garden, elev. 4000 ft, Kress 76-570 (DUKE), 78-901 (ou 78-1024 (DUKE). Panama. Bocas DEL Toro: N slope of Cerro Pate Macho, elev. 1200-1400 m, Andersson & Sytsma 1311 (s). Currigut: no further locality, Wagner s.n., April 1858 (mM); below Cerro Hornito, elev. 1000 m, Kress et al. 82-1368 (DUKE); Cerro Colorado, along road to copper mine beyond bridge over Rio San Félix near town of San Félix, elev. 1390 m, Croat 37288 (mo); Cerro Colorado, near continental divide, elev. 1500 m, Antonio 1494 (DUKE, MO), 1531 (DUKE, Mo), 1532 (Mo), 4859 (DUKE, MO), 4861 (MO), 4863 (Mo), 4912 (DUKE, MO); along road between Gualaca and Fortuna Dam site, 10.1 km NW of Los Planes de Hornito, elev. 1260 m, Antonio 4165 (mo); Gualaca—Fortuna Rd., elev. 1200-1400 m, Andersson 1306 (GB, s); NO de Campamento Fortuna (Hornito), sitio de regresando al SO del Campamento (Finca Pitti), elev. 1000-1200 m, Correa et al. s.n. (Mo); La Fortuna 1984] KRESS, HELICONIA 479 hydroelectric project, elev. ca. 1040 m, Hammel 1990 (mo); La Fortuna Dam site, elev. 1100 m, Kress & Hammel 83-1606 (DUKE); Kress et al. 82-1344 (DUKE), 82-1345 (DUKE), 82-1355 (DUKE), 82-1356 (DUKE); area E of main camp at Fortuna Dam site, elev. 1400- 1500 m, Folsom et al. 5447 (mo); 2 km en el camino Cerro de la Muerte, elev. 6000 ft, Correa 1244 (pMA); along road | km beyond Finca Alto Quiel toward Bajo Mono, elev. 5200 ft, Luteyn 3716 (DUKE, Mo); Cerro Horqueta, TTC-BMI Cloud Forest Litter Study, elev. ca. 1500 m, Duke et al. 13678 (DUKE, MO, SCz), 13667 (Mo); SW slopes of Cerro Horqueta ca. 6 km NW of Boquete, elev. 1700-1800 m, Wilbur et al. 15456 (DUKE); WwW Rio Caldera, elev. 4500-6500 ft, A/len 4656 (GH); Boquete, Palo Alto, elev. 1300-1772 m, Beliz 171 (pMA); Boquete, Finca Collins, Blum & Dwyer 2583 (mo, scz); Monte Rey near Boquete, Croat 15862 (mo); between Boquete and Monte Rey, Croat & Porter 15653 (F, Mo); E of Boquete on Cerro Azul near Quebrada Jaramillo, elev. 1500-1620 m, Croat 26791 (Mo); Boquete, elev. 3800 ft, Davidson 668 (F, GH, US); Boquete, 5 mi rom town near Finca Lerida, elev. 5300 ft, Kress et al. 77-822 (DUKE); pastures around El Boquete, elev. 1000-1300 m, Pittier 2936 (us); Bambito—Cerro Punta, elev. 2100 m, Camino entre Bambito y La Amenaza, elev. 6000 ft, Correa 1281] (PMA); roadside from Cerro Punta to Bambito, Lazor & Correa 2730 (Mo, scz); Bambito, elev. 1700 m, Andersson 1300 (s); lower slope of Baru, E of Bajo Choro region, elev. 6000-6500 ft, Hammel 3018 (Mo); above hwy. along Rio Chiriqui between Cerro Punta and Bambito, elev. 5200 ft, Kress & Clarkson 79-1088 (DUKE); in forest along Rio Chiriqui Viejo below Cerro Punta, elev. 5200 ft, Kress & Clarkson 79-1089a (DUKE), 79-1089b (DUKE); Rio Chiriqui Viejo, vic. of Nueva Suiza, elev. 5000 ft, Allen 6255 (us); on road from Rio Sereno to Volcan, Antonio 1304 (Mo); 2.2 mi E of El Hato del Volcan, Luteyn 802 (DUKE); Cerro Pate Macho, elev. 1500-1700 m, Andersson & Sytsma 1310 (GB, s); Rio Chevo, elev. 1650 m, Knapp 1451 (mo); Cerro Pelota, elev. 2300 m, Knapp 1499 (Mo). VE- RAGUAS: NE slopes of Cerro Delgadito just NW of Cerro Tute, S of town of Santa Fé, elev. 1000 m, Luteyn 4039 (DUKE); Cerro Tute, elev. 800-1400 m, Kress & Knapp &2- 1410 (DUKE). PHENOLOGY. Flowering principally February to August. HABITAT AND DISTRIBUTION. This species is restricted to altitudes above 1000 m on the Pacific slopes of the western highlands of Panama and neighboring Costa Rica (Map 4). Heliconia nutans inhabits disturbed primary and second- ary forests often near streams or rivers. It is the only Central American Hel- iconia with a pendent inflorescence that occurs at elevations over 2000 m. VARIATION. Heliconia nutans is composed of two morphologically distinct entities. One, wise occurs primarily around Boquete and Cerro Punta in Chiriqui, Panama, has persistent, papery floral bracts, distichous cincinnal bracts, entirely ee leaf blades, and a diminutive stature. The second entity is found in the mountains around La Fortuna and Cerro Colorado in eastern Chiriqui and has decomposing floral bracts, spirally arranged, somewhat con- torted cincinnal bracts, maroon or green leaf-blade undersides, and a more robust stature. Their distribution is not entirely allopatric, and where the two entities overlap (e.g., above Gualaca) the first is found at lower elevations and in drier habitats than the second. RELATIONSHIPS. Heliconia nutans is closely related to H. secunda, of Costa Rica 480 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 T T T T 1 J I L 1 Ap 4. Costa Rica and Panama, showing distribution of Heliconia nutans (squares) and H. platystachys (circles). and Nicaragua, with which it is often confused in the herbarium. The former species has a puberulous to tomentose rachis, a perianth 4—5 cm in length, and (in some cases) persistent floral bracts that appear papery when dried. 9. Heliconia platystachys Baker, Ann. Bot. (London) 7: 194. 1893 (excl. spec. cit. Donnell-Smith 1873 (kK), which is H. collinsiana Griggs var. velutina Kress). Tyre: [Colombia,] Santa Marta, Purdie s.n. (lectotype, Kk, chosen by Daniels, Kew Bull. 32: 682. 1978). PLATE IV, A, C. Bihai platystachys (Baker) Griggs, Bull. Torrey Bot. Club 31: 445. 1904. Heliconia catheta R. R. Smith, Phytologia 30: 65. 1975. Type: Panama, [Panama,] near old Fort pun mouth of Rio Chagres, Piper 6037 (holotype, us!). Herb with Musa-like habit, 3-4 m tall. Leafy shoots in groups of 3 to 20; pseudostem green, glaucous, 1.3—2 m tall, 4-5 cm in diameter; leaves 3 to 6 per shoot; petiole green, glaucous, 90-100 cm long, 1—1.3 cm in diameter; blade with the base unequal, cordate, the apex acute, the upper surface dark green, glabrous, with midrib green and glabrous, the lower surface green, usually glaucous, with midrib light green and glaucous, the longest blade |.3-1.7 m by 35-40 cm. Inflorescence pendent, to 80 cm long; peduncle red to green, pu- berulous to tomentose, 10-25 cm long, 1-1.6 cm in diameter; rachis flexuose, red becoming yellow near terminal cincinnal bracts, tomentose, 8-11 mm in diameter at base; cincinnal bracts spirally arranged, 10 to 20 per inflorescence, oriented 90—95° to axis of inflorescence, the basal bract sometimes sterile, the middle bract with apex acuminate, margins straight, inner surface puberulous and yellow-orange becoming green along margins, outer surface puberulous and red basally becoming yellow-green along margins and at apex, 11-16 cm long, 7.5—11 cm wide at base, I/w = 1.5. Floral bracts semitranslucent, decom- 1984] KRESS, HELICONIA 481 posing after anthesis, yellow, glabrous to minutely puberulous abaxially, 3.5— 4.5 cm long, 1-1.5 cm wide at base. Flowers 10 to 15 per cincinnus; pedicel pale yellow, puberulous, 1.5-2.5 cm long; perianth yellow to yellow-green, glabrous to minutely puberulous, 5.5-6 cm long, 1.1-1.3 cm wide at base, at anthesis curved 25-35°; free sepal reflexed, fused sepals with apices slightly reflexed; staminode white, 6-10 by 2-3 mm, apex apiculate; stamens with anthers flared outside perianth apex; pollen nutans type, large (62 x 76 um), suboblate (P/E = 0.82), subisopolar (DR/PR = 2.65), with the distal hemi- sphere constricted, convex, spinulose, the proximal hemisphere planar to sub- convex, psilate to minutely spinulose, the germination aperture distinctly pro- truding; ovary pale yellow, puberulous to tomentose, 7-10 by 7-8 mm. Drupes sparsely puberulous, 1.7-1.8 by 1.4-1.5 cm SPECIMENS EXAMINED. Costa Rica. PUNTARENAS: along hwy. 18 km N of Parrita, elev. sea level, Kress et al. 77-803 (DUKE). Panama. CANAL Zone: vic. of Nuevo Emperador, Blum 2394 (scz); near Fort Sherman Military Reservation, Maas & Mori 1754 (F, u); secondary tropical moist forest along road K-10, 4 km N of Arraijan, elev. 115 m, Nee 7160 (GH, PMA); near summit of Cerro Pelado, 1 km N of Gamboa, elev. 200-220 m, Nee 7439 (GH, MO, PMA); Rio Agua Salud, near Frijoles, Piper 5856 (us), Barro Colorado Is., NE of Barrunga Pt. at edge of lake, Croat 5616 (Mo), 5670 (scz); near #8 light house clearing, Croat 6387 (mo), Dodge 3486 (Gu), Fairchild 17 (F), Kenozer 232 (us), Summit Garden, Croat 10790 (F, Ny); Albrook Air Force Base Research Forest Site, Stimson 5068 (DUKE, PMA, Scz); Albrook, Dwyer 6599 (mo); along Pipeline Rd. near Gamboa elev. ca. 75 m, Kress & Cooper-Smith 76-643 (DUKE), Kress et al. 77-860 (DUKE); Gaillard way to Nombre de Dios, elev. 150-300 m, Croat 26113 (mo); N of Maria Chiquita on road to Portobelo, Croat 11353 (Mo), Knapp & Mallet 5718 (mo), between Gatun Lock and Fort Sherman, ca. sea level, Kress 80-1243 (DUKE), along Hwy. 79, Quebrada Sar- dinilla, 6 km NE of Buena Vista, elev. 55 m, Nee 657] (Mo). DARIEN: Punta Guayabo Grande, along beach, elev. 0-50 m, Antonio & Hahn 4217 (mo), Knapp & Mallet 3036 (mo); Santa Fé, elev. ca. 15 m, Duke 14269 (mo); vic. of Paya, Rio Paya, Stern et al. 427 (GH, MO, UC, Us); SW of Jaqué, Sytsma & D/Arcy 3483 (mo); vic. of Campamento MO, UC, US). PANAMA: Vacamonte Pt., Allen 2959 (GH, Mo); Cerro Campana, elev. 600 m, yee 1260 (mo); vic. of Rio Tapia, Bartlett & Lasser 16623 (mo); TTC Albrook Hwy. on road to Cerro Azul, Croat 11512 (scz), near old Fort Lorenzo, mouth of Rio Chagres, Piper 6037 (us); Agricultural Exp. Station at Matias Hernandez, Pittier 6813 (us); near big swamp E of Rio Tocumén, Standley 26726 (us), 5 mi W of Chepo near Interamerican Hwy., Tyson 6701 (Mo, PMA). SAN Bias: Playén Chico, Stier 1] (Mo), 130 (Mo, US). PROVINCE UNKNOWN: Kuntze $.n., Aug. 1874 (Ny). PHENOLOGY. Flowering primarily June to October. HABITAT AND DISTRIBUTION. This species occurs in undisturbed forest and open secondary growth usually at elevations of less than 100 m, although several collections have been made at 600 m. Heliconia platystachys is distributed sporadically along the southern Pacific coast of Costa Rica and into Panama, 482 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 where it becomes more abundant in the provinces of Panama, Col6n, and Darién (Map 4). It is also found in northern South America. RELATIONSHIPS, In Central America, Heliconia platystachys, a member of the nutans group, 1s closely related to H. collinsiana, but it has closer relatives in South America. The puberulous cincinnal bracts and ovaries readily distinguish H. platystachys from other species of the nutans group. NOMENCLATURE. The confusion over typification of Heliconia platystachys has been addressed by Daniels (1978). The problem arose when H. platystachys was redescribed by Smith (1975) as H. catheta from Panama. Smith made this decision after seeing only part of the type collection of H. platystachys. The type housed at Kew consisted of two separate entities: a Colombian collection made by Purdie and a Guatemalan one made by Donnell-Smith. The latter collection, the one seen by Smith, is attributable to H. collinsiana. Daniels, based on Baker’s protologue, accurately chose the Purdie collection as the lectotype of H. platystachys, thereby reducing Smith’s H. catheta to synonymy. 10. Heliconia collinsiana Griggs, Bull. Torrey Bot. Club 30: 648. 1903. Type: Guatemala, Alta Verapaz, near Finca Sepacuité, Cook & Griggs 352 (us!) Bihai collinsiana (Griggs) Griggs, Bull. Torrey Bot. Club 31: 445. 1904. Heliconia rostrata auct. non Ruiz & Pavon: Standley & Steyerm. Fieldiana Bot. 24: 2 Herb with Musa-like habit, 4-6 m tall. Leafy shoots in groups of 3 to 50: pseudostem glabrous to tomentose (sometimes glaucous), 1.2-3 m tall, 3.5-7 cm in diameter; leaves 3 to 5 per shoot; petiole green, glaucous, 40-120 cm long, 1.3—3 cm in diameter; blade with the base unequal, obtuse, the apex acute, the upper surface green to dark green, glabrous, with midrib green and glabrous, the lower surface green, glabrous or with white waxy coating, midrib light green, or maroon and glaucous, the longest blade 1.1-2.5 m by 28-70 cm. Inflorescence pendent, to 72 cm long; peduncle green to red, glabrous to ve- lutinous with rusty hairs, 10-34 cm long, !|.1-1.9 em in diameter; rachis flex- uose, red (sometimes becoming yellow near apex), puberulous to velutinous, 6-13 mm in diameter at base; cincinnal bracts spirally arranged, 7 to 18 per inflorescence, oriented 90-100° to axis of inflorescence, the basal bract some- times sterile, the middle bract with apex acuminate, margins straight to rev- olute, inner surface yellow-pink to red and glabrous, outer surface red to orange- red and glabrous to glaucous, 11-17 cm long, 6-8 cm wide at base, \/w = 2.0. Floral bracts opaque and decomposing after anthesis, white to pale yellow, glabrous to puberulous abaxially, 4.5-7 cm long, 1.5-2.5 cm wide at base. Flowers 10 to 20 per cincinnus; pedicel yellow to orange, glabrous to puberulous, 1-3 cm long; perianth yellow to orange, reddish along sepal mar- gins, glabrous to puberulous, 4.6-6 cm long, 8-11 mm wide at base, at anthesis curved 15-25% free sepal sometimes reflexed, fused sepals with apices not reflexed; staminode white, apex 1-1.7 cm by 3-6 mm, tridentate with center tooth longer than laterals; stamens with anthers flared outside perianth apex; 1984] KRESS, HELICONIA 483 pollen nutans type, small to medium size (52-60 x 52-65 um), oblate-sphe- roidal (P/E = 0.92-0.99), subisopolar (DR/PR = 2.67-3.28), with the distal hemisphere convex, spinulose, the proximal hemisphere subconvex, psilate to verrucose, the germination aperture distinctly protruding; ovary yellow to pale green, glabrous, 6-10 by 6-10 mm. Drupes glabrous, 1.2-1.5 by 1.6-1.8 cm. DIstTRIBUTION. This species, found from Mexico to Nicaragua (Map 5), is com- posed of two varieties that may overlap in geographic distribution but are separated by elevational differences. ReLATIonsHIPs. Heliconia collinsiana and H. platystachys, both members of the nutans group, are more closely related to each other than to any other Central American members of the genus. However, each of these taxa may be more closely related to some South American species than it is to the other. Heliconia collinsiana can be distinguished from H. platystachys by its pure red cincinnal bracts and its glabrous floral bracts and ovaries. REMARKS. Several collections of a taxon that appears to be very closely related to Heliconia collinsiana var. velutina have been made in the states of Guerrero d Nayarit, Mexico. These plants differ from var. ve/utina in their much smaller stature (ca. 2 m) and their pubescent to tomentose calyxes. Smith (1968) named these individuals H. x mooreana, believing them to be hybrids of H. collinsiana and H. schiedeana. Because of the rarity of natural hybrids in Heliconia (see section on hybrids) and the multiple collections of this taxon from several different localities in Mexico, Smith’s interpretation is not ac- cepted here. However, a decision as to the varietal status of this taxon must await further collections. 9 =] SPECIMENS EXAMINED. Mexico. GUERRERO: near km 339 on hwy. to Acapulco below Acahuizatla, elev. 940 m, Moore 6204 (GH); Acapulco and vic., Palmer 311 (GH, K, US). Nayarit: Arroyo de la Cosiscadora, elev. 60 m, Ortega 49 (us). Key to the Varieties of Heliconia collinsiana Leaf blades with lower surface glaucous; peduncle and rachis cane to tomentose; Hanth and Ovary yellows 250645) Kenta eaeiee shave eee 10a. var. collinsiana. Leaf blades with lower surfacen not ee peduncle and rachi eet e velutinous; perianth orange; Ovary green. .... 0.6... eee eee 10b. var. velutina. 10a. Heliconia collinsiana Griggs var. collinsiana Pate IV, B, D. Herb to 4 m tall. Leafy shoots in groups of 5 to 50; pseudostem glaucous, 1.2-1.8 m tall, 3.5-5 cm in diameter; leaves with the petiole light green, 40- 62 cm long, 1.3-1.5 cm in diameter, the blade having lower surface green with white waxy coating, the longest blade 1.1-1.5 m by 28-43 cm. Inflorescence with peduncle green to red, glabrous to tomentose, 10-34 cm long, 1.1-1.4 cm in diameter; rachis red, yellow near apical cincinnal bracts, puberulous to tomentose, 6-10 mm in diameter at base; cincinnal bracts 7 to 14 per inflo- rescence, the basal bract sometimes sterile, the middle bract with margins 484 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 i} T T T T Mexico and Central America, showing distribution of Heliconia collinsiana vars. es (circles) and velutina (triangles). straight, inner surface yellow-pink to red, outer surface red to orange-red and often glaucous, 11-17 cm long, 6-8 cm wide at base. Floral bracts white to pale yellow, glabrous abaxially, 4.5—-7 cm long, 1.5-2 cm wide at base. Flowers 10 to 12 per cincinnus; pedicel yellow to orange-yellow, glabrous to puberulous, 1.3-3 cm long; perianth yellow to orange-yellow, glabrous, 4.6-6 cm long, 8- 11 mm wide at base, at anthesis curved 15-25°; staminode 1.1-1.7 cm by 3- 6 mm wide; ovary yellow, 6-10 by 6-10 mm. SPECIMENS EXAMINED. Mexico. CuH1apas: NW side of Cerro Vernal, 25-30 km SE of Tonala, elev. 400-600 m, Breedlove 25595 (DUKE, Mo); 4 mi N of Tapachula along road to Nuevo Aleman, elev. 250 m, Croat 43790 (Mo); 8.5 mi NE of Escuintla on gravel road to El Triunfo, elev. 250 m, Croat 43825 (mo); between Esquintla and Monte Ovando, .8 km NW of Turquiz, elev. 100 m, Croat 47469 (mo); Esperanza, Escuintla, Matuda 16693 a 18019 (F); Aguacorte Palenque, Matuda 3823 (Gu); road to microwave tower about mi from Hwy. 200, hwy. entrance 5.5 mi NW of Tres Picos, elev. 200 m, anne - al. 2412 (Gu); Finca Prusia, municipio de Angel Albino Corzo, elev. 2400 ft, Ton 359] (puKeE),; N of Huixtla, elev. 500 m, Taylor see (DUKE). shee atin aera Sierra Madre, elev. 5-800 m, Anonymous s.n., Jan. 1899 (Gu, us). NAYARIT: 8-10 m (DUKE). OAXACA: Distr. of Tuxtepec, Chiltepec and vic., elev. 20 m, Martinez-Calder6n 99 (GH, US); ravine 35 km E of Mathias Romero, 4 km W of Palomares on trans-isthmus highway, Moore 8111 (a). VERACRUz: Fortin de las Flores (posada Loma), elev. 850 m, cultivado, Nevling & G6mez-Pompa 312 (Gu). Guatemala. ALTA VERAPAZ: 22 mi from of secanquim, elev. 550 m, Maxon & Hay 3187 (us); near Eneaene, elev. 360 m, Standley ermark 31128 (F). ESQUINTLA: between Rodeo and Osuna, below Volcan de Fuego, elev. 2300 ft, Kress et al. 77-753 (DUKE); La Trinidad on road between Escuintia and Finca 1984] KRESS, HELICONIA 485 Zapote, elev. 780 m, Standley 65042 (F); Masagua, elev. 400 m, J. D. Smith s.n., April 1890 (Us). HUEHUETENANGO: between Ixcan and Rio Ixcan, Sierra de los Cuchumatanes, elev. 150-200 m, Stevermark 49342 (F). IZABAL: 10-15 km W of El Estor, thickets along road in flat terrain on red clay, Harmon & Dwyer 4359 (mo, us); vic. of Quirigua, elev. 75-225 m, Standley 24480 (GH, US). QUEZALTENANGO: El Palmar, elev. 2300 ft, Keller- man 6078 (F); Palmar, elev. 2300 ft, Kellerman s.n., Feb. 1906 (us). RETALHULEU: Rio Talculan W of Retalhuleu, elev. 300 m, Standley 87338 (F). SAN Marcos: Rio below Rodeo, elev. 600 m, Standley 68796 (F). SANTA Rosa: near El Molino, elev. 600 m, Standley 78520 (F); region of Platanares, between Taxisco and Guazacapan, elev. 220 m, Standley 79130 (F). SUCHITEPEQUEZ: Rio Sis, elev. 1300 ft, Donnell-Smith 2804 (K, Us); along Rio Madre Vieja, above Patulul, elev. 450 m, Standley 62244 (F); E slopes of Volcan Santa Clara, above Chicacao, elev. 1250 m, Steyermark 46777 (F); vic. of Tiquisate, elev. 100 m, Steyermark 47633 (F). Belize. STANN CREEK: Middlesex, elev. (us); Finca Santa Maria, W of Santa Tecla, elev. 750 m, Carlson 162 (F), elev. 800 m, Inst. Trop. Inv. Cient. (ITIC) 1206 (Mo). SANTA Ana: elev. 700 m, J77C 1201] (Mo). SAN SALVADOR: elev. 670 m, [TIC 1204 (Mo); vic. of San Salvador in thicket along stream, elev. 650-850 m, Standley 20551 (us). SAN VICENTE: vic. of San Vicente, elev. 350-500 (F); rain forest on slopes, Lancetilla Valley ca. 3 mi S of Tela, elev. 200-500 ft, Webster et al. 12642 (F, Mo); Lancetilla Valley, near Tela, Pfeifer 2126 (us); E of La Ceiba, in rocky ravine, slopes near Puerte Alto on S.F. Co. R.R., elev. 800 ft, Yuncker et al. 8537 (F, GH, MO, NY, S, UC, US). COLON: Trujillo, Rio Nesro, Saunders 556 (Ny), 703 (Ny). Cortes: Mt. Bella Vista above San Pedro Sula, Barkley & Leiva s.n., 4 Oct. 1969 (Gu). OLANCHO: Orillas del Patuca, cerca Rio Guayape, G. Cruz s.n., 2 March 1973 (mo); rich soil of montane forest between Cofradia and Rancho Quemado, Barkley & Errazuri 40307 (Mo). Nicaragua. GRANADA: Comarca La Fuente, elev. 600 m, Castro & Montiel 609 (DUKE, Mo); Volcan Mombacho, elev. 350-500 m, Moreno 413 (Mo), 439 (mo), 1466 (Mo). JINOTEGA: NE de Wiwili, entre El Carmen y Wamblan, elev. 250-400 m, Ara- quistain & Castro 1859 (DUKE), Araquistain & Moreno 1508 (DUKE). MANAGUA: 2 mi from El Crucero on the Pan Am Hwy., elev. 2700 ft, Kress et al. 77-758 (DUKE); between El Crucero and house of Finca Santa Julia, elev. 600-900 m, Standley 8380 (F); along 8 ca. 2.4 km SW of intersection with Hwy. 2, km 28, elev. ca. 700 m, Stevens 3982 (DUKE), 5167 (DUKE). NUEVA SeGovia: Jalapa, Atwood et al. 6773 (Mo). ZELAYA: tall forest about Yauya, 18 km SE of La Luz, elev. 150-200 m, Bunting & Licht 581 (Fr, NY). PHENOLOGY. Flowering mainly January to August (dry season and early part of rainy season); some mature inflorescences all year. HABITAT AND DISTRIBUTION. Variety co//insiana inhabits open secondary growth and well-drained soils at middle to higher elevations (up to 1200 m) from southern Mexico to central Nicaragua (Map 5). It is the northernmost species of Heliconia with a pendent inflorescence and is almost completely allopatric with all other species having the same inflorescence type. 10b. Heliconia collinsiana Griggs var. velutina Kress, J. Arnold Arbor. 69: 255. 1981. Type: Guatemala, Quezaltenango, lower slopes of Volcan Santa Maria de Jests, along hwy. to Quezaltenango near Santa Mari de Jestis, elev. 1450 m, Kress, Clarkson, & McDade 77-756 (holotype, DUKE; isotypes, F, K). 486 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Herb to 6 m tall. Leafy shoots in groups of 3 to 8; pseudostem glabrous to tomentose, 2.5—-3 m tall, 7 cm in diameter; leaves with the petiole green, 1- 1.2 m long, 2.5-3 cm in diameter, the blade with lower surface green and glabrous, the longest blade 2.2-2.5 m by 65-70 cm. Inflorescence with peduncle rusty red, velutinous to villous with rusty hairs, 16-30 cm long, 1.5-1.9 cm in diameter; rachis red, tomentose to velutinous, 1.1—-1.3 cm in diameter at base; cincinnal bracts 15 to 18 per inflorescence, the basal bract usually fertile, the middle bract with margins revolute, inner surface red, outer surface deep red and glabrous, 14-17 cm long, 6-7.5 cm wide at base. Floral bracts white, glabrous to puberulous abaxially, 5-6 cm long, 1.5—2.5 cm wide at base. Flowers 15 to 20 per cincinnus; pedicel pale orange, puberulous, 1-1.7 cm long; perianth pale orange, glabrous to puberulous, 5.5-5.7 cm long, 8-9 mm wide at base, at anthesis curved 15—20°; staminode 9-12 by 4-5 mm; ovary pale green, 7- 8 by 7-9 mm SPECIMENS EXAMINED. Guatemala. EscuINTLA: Finca Monterrey, S slope of Volcan de lower S-facing slopes of Volc4n Santa Maria between Santa Maria de Jesas and Cala- huache, elev. 1300-1400 m, Steyermark 33575 (F). SANTA Rosa: Cenaguilla, elev. 4000 ft, Heyde & Lux 4635 (GH, K, US). SUCHITEPEQUEZ: Finca Moca, in bushy growth at edge of barranca, elev. 5000 ft, Skutch 2101 (Gu). El Salvador. AHUACHAPAN: Sierra de Apaneca in region of Finca Colima, Standley 20081 (us). SANTA ANA: moist canyon slope, W side of Cerro del Aguila, elev. 1650 m, Tucker 1291] (F, K, UC, US), SONSONATE: Cerro los Naranjos, [TIC 1207 (mo). PHENOLOGY. Flowering and fruiting mainly during dry season (January to April) and early part of rainy period (May to July); some inflorescences produced all year. HABITAT AND DISTRIBUTION. Heliconia collinsiana var. velutina is found on steep forested slopes at middle to higher elevations (> 1300 m) in the Gua- temalan and Salvadorian highlands (Map 5). It is one of the few Central Amer- ican heliconias found at such high elevations. RELATIONSHIPS. This variety is distinguished from var. collinsiana by its greater stature, its lack of glaucous lower leaf surfaces, its hairy peduncle and rachis, its orange flowers, and its growth at higher altitudes (above 1300 m). 11. Heliconia curtispatha Petersen in Martius & Eichler, Fl. Brasil. 3: 15. 1890. Tyre: Panama, [Canal Zone,] prope Gatin, M. Jan. Flor., 1. Wagner 5.n., “Tan, 1858 (holotype, M!). Piate V, A, C. Bihaia curtispatha (Petersen) Kuntze, Rev. Gen. Pl. 2: 685. 1891. Bihai longa Griggs, Bull. Torrey Bot. Club 31: 446. 1904; Heliconia longa (Griggs) Winkler in Engler & Prantl, Nat. Pflanzenfam. ed. 2. 15a: 536. 1930. Type: Costa Rica, Alajuela, El Pez road from Buena Vista to San Carlos Valley, Cook & Collins 47 (holotype, us!). Heliconia rostrata auct. non Ruiz & Pavon: Woodson & Schery, Ann. Missouri Bot. 1984] KRESS, HELICONIA 487 Gard. 32: 54. 1945, in part (Woodson & Schery included H. curtispatha and a hybrid in H. rostrata). Herb with Musa-like habit, 6-7 m tall. Leafy shoots in groups of 5 to 25; pseudostem green, glabrous, 1.8-—3 m tall, 5—6.5 cm in diameter; leaves 4 to 7 per shoot; petiole green to reddish green, glabrous, 90-145 cm long, 1.5-2.3 cm in diameter; blade with the base unequal, cordate, the apex acute, the upper surface dark green, glabrous, with midrib light green to green and glabrous, the lower surface green, covered with white waxy coating, with midrib yellow to red-green and glabrous, the longest blade 1.9—2.3 m by 35-55 cm. Inflorescence pendent, to 1.6 m long; peduncle red (sometimes yellow), glabrous to scurfy, 28-37 cm long, 1.3—1.8 cm in diameter; rachis flexuose, red to red-orange puberulous to scurfy, |.2—1.8 cm in diameter at base; cincinnal bracts distichous to spirally arranged, 20 to 40 per inflorescence, oriented 130-145° to axis of inflorescence, the basal bract usually sterile, the middle bract with apex acute to acuminate, margins straight to involute, inner surface yellow-orange to pink and puberulous to tomentose with rusty hairs, outer surface red to yellow- orange near rachis and puberulous, 8.5—11.5 cm long, 6.5—9 cm wide at base, l/w = 1.3. Floral bracts opaque, persistent, white to yellow, tomentose to ve- lutinous abaxially, 5.5-6.5 cm long, 1.6-2.3 cm wide at base. Flowers 10 to 20 per cincinnus; pedicel white to pale yellow, glabrous to puberulous, 1.1-2 cm long; perianth white to yellow and glabrous basally, deep yellow and slightly puberulous toward apex, 5.5—6.5 cm long, 9-10 mm wide at base, at anthesis curved 70—90° and parabolic; free sepal somewhat reflexed, fused sepals with medium size (57 x 71 wm), suboblate (P/E = 0.82), isopolar (DR/PR = 0.99), with the distal hemisphere subconvex, spinulose, the proximal hemisphere convex, minutely spinulose, the germination pore distinct and somewhat sunk- en; ovary white, glabrous, 7-11 by 5-7 mm. Drupes glabrous, 1.7 by | cm. SPECIMENS EXAMINED. Nicaragua. ZELAYA: 5 mi W of Bonanza, Atwood & Neill 6995 (Mo); 20 mi W of Rama near Rio Micito, elev. 200 ft, Kress et al. 77-759 (DUKE), vic. of El Recreo, on Rio Mico, elev. ca. 30 m, Standley 19407 (F). Costa Rica. ALAJUELA: 5 km S of Canalete near Rio Zapate and along new road to Upala, elev. 100—200 m, Burger & Baker 10000 (F); upper Rio Sarapiqui near Cariblanco and along road to Colonia Virgen del Socorro, elev. 800 m, Burger & Antonio 11133 (F); steep bank above Quebrada Arrayanes, Cariblanco, elev. 900 m, Lent 3536 (F); 8 km NE of Villa Quesada, near Artezalea and Methodist Rural Center, elev. 550 m, Molina et al. 17219 (F, Ny); forests along Rio San Rafael, near hot springs, Hacienda La Marina, ca. 14 km NE of Villa Quesada, elev. 50 m, Molina et al. 17422 (F, Ny); near Santa Clara, elev. 2100 ft, Kress et al. 77-768 (DUKE); near San Miguel, elev. 1700 ft, Kress & Cooper-Smith 76-531 (DUKE); near San Miguel, Sheffy 69 (cr). CARTAGO: near Turrialba, 2 km W of Oriente along Rio Vueltas, Sheffy & Tosi 58 (cr). HEREDIA: Finca La Selva, new property, Hammel 11459 (DUKE). LIMON: between Siquirres and Rio Pacuare, and hills S of RR bridge over Rio Pacuare, elev. 50-100 m, Burger & Liesner 7000 (F); in old banana plantation 1 km W of Moin, elev. 25 m, Kress et al. 76-617 (DUKE); 5 mi from Puerto Limon on road to Siquirres, elev. ca. 100 ft, Kress et al. 77-789 (DUKE); Cimarrones, ca. 10 km E of Siquirres, elev. 200 m, Maas 1133 (F, U). PUNTARENAS: San Vito de Java, cult. at Las Cruces Trop. Bot. Garden, elev. 4000 ft, Kress 76-577 (DUKE), 78-1023 (DUKE). SAN José: bords du rio de las Vueltas, Tucurrique, elev. 635 m, Ténduz 12923 488 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 (K, M, US, W). PROVINCE UNKNOWN: foréts de Tsallo, elev. 200 m, T6nduz 9433 (us, W); lieux humides a Zent, elev. 31 m, Ténduz 14542 (F (photo), NY). Panama. CANAL ZONE: elev. 1600-1700 ft, Kress et al. 80-1165 Vise ie He Cenone er elev. 700-800 m, Andersson & Sytsma 1255 (Mo, $). COLON: fore hill d 18 km past Sardinilla on way to Nombre de Dios, elev. 150-300 m, Ca 26093 ane Cerro Santa Rita, elev. 800-900 ft, Antonio 1787 (mo); Santa Rita Ridge, Dwyer & Gentry 9549 (F, MO), Sytsma 1583 (mo); between France Field, Canal Zone, and Cartival, Standley 30425 (us); along Rio Guanche, elev. 40 m, Antonio 1217 (mo); Rio Guanche, elev. 50 ft, Kress & Hammel 1241 (G, Mo, Ss). DARtéN: Punta Guayabo Grande to Rio Jaqué, elev. 5-200 m, Antonio & Hahn 4384 (mo); Ensefiada del Guayabo, Garwood 1200 (mo); between Quebrada Venado and Peje Swamp on headwaters of Rio Tuqueza, Bristan 1041 (mo); E slope of Cerro Sapo, elev. 2500 ft, Hammel 1319 (mo); Cerro Pirre, Bristan 517 (NY). PANAMA: 3 mi above Goofy Lake near Cerro Azul, Croat 11571 (F, Mo, Ny); Cerro Jefe, D'Arcy et al. 1957 (F, Mo), elev. 2300 ft, Kress et al. 77-864 (puke); Alto de Pacora, below Cerro Jefe, elev. 800-900 m, Kress et al. 76-635 (puKE), Kress & Hammel 83-1573 (DUKE); Campo Tres, 3 mi NE of Alto de Pacora, elev. 500-800 m, Liesner 537 (mo); road to Carti, Antonio 1288 (mo); El Llano—Carti Rd., elev. 100-200 m, Andersson & Sytsma 1259 (GB, MO, S$), elev. 300-400 m, Folsom 2561 (mo), Folsom et al. 6190 (Mo), Kress & Hammel 82-1333 (DUKE); 6 km nbove Pan Am Hwy. on road from El Llano to Carti- Tupile, elev. ca. 200 m, Kennedy 1774 (F, Mo, Ny); 2.4 mi N of Interamerican Hwy. toward El Llano and Carti, elev. ca. 1700 ft, Luteyn 4 oa (DUKE); El Llano—Carti Rd., 5 km N of Pan Am Hwy. at El Llano, elev. ca. 300 m, Nee 7928 (Mo, scz), premontane wet forest, Nee 7944 (Mo, Us). SAN BLAs: hills SW of Puerto Obaldia, Croat 16721 (mo). PHENOLOGY. Flowering throughout year, with slight peak during early rainy season (June to September) HABITAT AND DISTRIBUTION. At higher elevations (above 100 m), Heliconia curtispatha inhabits old secondary growth and undisturbed forested slopes. At lower elevations, especially around the Lim6n area in Costa Rica, this species will invade old banana plantations and other disturbed areas. Heliconia cur- tispatha is distributed from southern Nicaragua to eastern Panama and Co- lombia. It is primarily found on the Caribbean side of the central cordilleras of Costa Rica and Panama from sea level to 900 m (Map 6). VaRIATION. Heliconia curtispatha is generally morphologically homogeneous except for a distinct color form that is found in the Caribbean lowlands around Limon in Costa Rica. Plants found in this area have distinctive orange to yellow rachises, as opposed to the normal red ones common throughout the range RELATIONSHIPS. This species shares a unique common ancestor with Heliconia stilesi, together these species constitute the monophyletic curtispatha group. his group is allied to the pogonantha group but differs in its leaf blades with glaucous lower surfaces and its glabrous, parabolic perianths, as well as in several pollen characters. NoMENCLATURE. Daniels and Stiles (1979) mistakenly applied Griggs’s syn- onym Heliconia longa to this taxon. 1984] KRESS, HELICONIA 489 iv T q J T ee e e Pad & 5° e 104 v e Vv e e e v e v vv Ce e v . © ye — ca e e@ 4 e gq” x 7 86 7 i | is Map 6. uthern Nicaragua, Costa Rica, and Panama, showing allopatric distribution of Heliconia curtispatha (circles) and H. stilesii (triangles). 12. Heliconia stilesii Kress, Brenesia 19/20: 202. 1982. Type: Costa Rica, Puntarenas, slopes above airport at Golfito, elev. 50 ft, Kress & Clarkson 79-1096 (holotype, DUKE!; isotypes, cR!, F!). FiGuRE 11; PLATE V, B, D. Heliconia curtispatha auct. non Petersen: ce Bull. Torrey Bot. Club 42: 316. 1915; Daniels & Stiles, Brenesia 15(Supl.): 19. 1979, Heliconia platystachys auct. non Baker: sien Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 184. 1937. Herb with Musa-like habit, 5-6 m tall. Leafy shoots in groups of 7 to 16; pseudostem pale yellow-green, glabrous to slightly glaucous, 2.4—3.5 m tall, 6- 7 cm in diameter; leaves 4 to 6 per shoot; petiole yellow-green, glabrous, |.2- 1.9 m long, 1.5-2.3 cm in diameter; blade with the base unequal, truncate to cordate, the apex acute, the upper surface dark green, glabrous, midrib light green and glabrous, the lower surface light green, covered with white waxy coating, midrib yellow-green with maroon and glabrous, the longest blade 1.7- 2.3 m by 45-55 cm. Inflorescence pendent, to 1 m long; peduncle red, glabrous to scurfy, 10-75 cm long, 1.1—1.8 cm in diameter; rachis strongly flexuose with cincinnal bracts often touching each other, red, scurfy, 1-1.4 cm in diameter at base; cincinnal bracts distichous, 20 to 35 per inflorescence, oriented 135- 155° to axis of inflorescence, the basal bract often sterile, the middle bract with apex acute to acuminate, margins straight to overlapping near rachis, inner surface pink to white and pubescent to tomentose with rusty hairs, outer surface rose-red and puberulous to scurfy, 8.5—9 cm long, 6-7.5 cm wide at base, I/w = 1.3. Floral bracts opaque, persistent, pale yellow (sometimes pink), tomentose to velutinous abaxially, 4.5-6.5 cm long, 1.4-1.8 cm wide at base. Flowers 10 to 20 per cincinnus; pedicel white to yellow, glabrous, 1.5—1.9 cm long; perianth white to pale yellow basally, deep yellow toward apex, glabrous, puberulous 490 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 along margins, 5.1-6.2 cm long, | cm wide at base, at anthesis curved 55-80° and parabolic; free sepal straight to slightly reflexed, fused sepals with apices slightly reflexed; staminode white, 5-8 by 3-4 mm, apex cuspidate; stamens with anthers connivent inside apex of corolla tube; pollen curtispatha type, medium size (53 x 66 um), suboblate (P/E = 0.82), isopolar (DR/PR = 1.17), with the distal hemisphere convex to subconvex, spinulose, the proximal hemi- sphere convex, minutely spinulose, the germination aperture distinct and some- what sunken; ovary white, glabrous, 8-9 by 5-6 mm. Drupes glabrous, 1.2- 1.5 cm by 7-9 mm. SPECIMENS EXAMINED. Costa Rica. PUNTARENAS: 5 km W of Palmar Norte on road to Puerto Cortes, elev. 300 m, Lent 158 (cr, F); Esquinas Forest, elev. 30 m, Allen 6289 (GH); entre Los Rios Piedras Blancas y Esquinas, Osa, elev. 75 m, Jiménez 2249 (F, Ny); Esquinas Forest Reserve near Rio Claro, elev. ca. 100 ft, Kress 78-1047 (DUKE); slopes above Golfito along trail to TV tower, elev. 100- 300 m, Burger & Matta 4769 (F, Gu), 4771 (F, us); 12 km E of Golfito, Harmon & Fuertes 6173 (mo); Golfito, near airport, elev. ca. 50 ft, Kress 78-1048 (DUKE); 3.5 mi beyond Golfito, ca. sea level, Kress et al. 76-586 (DUKE); 5 mi above Villa Neily on road to San Vito de Java, elev. 2100 ft, Kress 78-1050 (DUKE); San Vito de Java, cult. at Las Cruces Trop. Bot. Garden, elev. 3900 ft, Kress 76-576 (DUKE), 78-1032 (DUKE); (Onillas de Mar de Golfito de Osa, Brenes 12325a m (F), Burger & Stolze 5469 (cr, F); 4 mi W of Rincén de Osa, near airport, elev. 100 ft, Raven 21686 (cr, F, NY); Rincén, Semple 80 (Mo), Shep 37 (cr), SAN JOSE/PUNTARENAS: between Playa Dominical, Baru, and Tinamastes along road to San Isidro del General, elev. 10-500 m, Burger & Baker 10138 (F). SAN José: along hwy. between Parrita and Santiago near Finca Los Angeles, Kress & Cooper-Smith 76-634 (DUKE), elev. 2200 ft, Kress et al. 77-799 (DUKE). Panama. CuHiriQui: Peninsula de Burica, 10 mi W of Puerto Armuelles, elev. 100-300 m, Liesner 85 (F, MO, NY). PHENOLOGY. Flowering mainly during early part of rainy season (June to Sep- tember); some inflorescences produced all year. HABITAT AND DISTRIBUTION. This species is often found on steep slopes in old secondary growth at elevations below 500 m. It is primarily restricted to the drier Pacific slopes and coastal plains of Costa Rica, from the area around Parrita south to the Burica Peninsula (Map 6). ReLationsuips. Heliconia stilesii is most closely related to H. curtispatha, the only other Central American member of the curtispatha group. It differs from that species in its extremely flexuose rachis, in its strictly distichous, nearly imbricate cincinnal bracts, and in being mainly restricted to the Pacific slopes of Costa Rica. Figure 11. Heliconia stiles. A, inflorescence. B, C, cincinnal bracts: B, entire (Ba, stigma. (© Museo Nacional de Costa Rica, 1982; reprinted with permission.) KRESS, HELICONIA 491 1984] Sy “ >>> iN ae 492 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 NoMENCLATURE. Griggs (1915) and later Daniels & Stiles (1979) mistakenly applied the name Heliconia curtispatha Peterson to this taxon. 13. Heliconia sessilis Kress, J. Arnold Arbor. 62: 251. 1981. Type: Panama, Col6n, along road between Gatin Lock and Fort Sherman, elev. ca. 3 m, Kress 80-1241 (holotype, DUKE!; isotypes, F!, GH!, K! Mo!, PMA!). FiGure 12; PLate VI, A, C. Heliconia pogonantha auct. non Cuf.: Croat, Fl. Barro Colorado Is. 257. 1978. Herb with Musa-like habit, 6-7 m tall. Leafy shoots in groups of 10 to 20; pseudostem green to brown, glabrous, 2.5 m tall, 8-9 cm in diameter; leaves 4 or 5 per shoot; petiole olive green, glabrous, 1.4—1.8 m long, 2 cm in diameter; blade with the base unequal, attenuate to obtuse, the apex acute, the upper surface green, glabrous, with midrib olive green and glabrous, the lower surface dull green, glaucous, with midrib yellow-green with central maroon stripe and slightly glaucous, the longest blade 2 m by 40-60 cm. Inflorescence nodding to pendent, to 90 cm long; peduncle obscure, red, puberulous, 2 cm in diameter, rachis flexuose, red at base of inflorescence, yellow toward distal bracts, pu- berulous, 1.8-1.9 cm in diameter at base; cincinnal bracts spirally arranged, 23 to 30 per inflorescence, oriented 90° to axis of inflorescence, the basal bract usually sterile, the middle bract with apex acute, margins straight becoming involute near rachis, inner surface white to pink and glabrous to puberulous, the outer surface red and glabrous, 8-9 cm long, 9 cm wide at base, I/w = 0.9. Floral bracts opaque, persistent, pale yellow, pink along margins, puberulous abaxially, 3.5-4 cm long, 1.8-2.1 cm wide at base. Flowers 15 to 20 per cincinnus; pedicel ie reddish near ovary, glabrous, o- 10 ee long; ela deep yellow distally, paler at base, glab except for argins 4.8-5.1 cm long, 1.1—1.2 cm wide at base, at anthesis curved 30- 40°; and ‘slightly sigmoid; free sepal reflexed, fused sepals with apices reflexed; staminode white to pale yellow, 6-7 by 3 mm, apex cuspidate; stamens with anthers connivent inside apex of corolla tube; pollen medium size (57 x 79 um), euoblate (P/E = 0.72), subisopolar (DR/PR = 2.43), with the distal hemisphere sub- convex, spinulose, the proximal hemisphere planar, psilate, the germination aperture distinct, flattened, sunken; ovary white becoming pale lavender around apex, glabrous, 1-1.2 cm by 6-7 mm. No mature fruits seen. SPECIMENS EXAMINED. Panama. CANAL ZONE: vic. of Fort Sherman, W of Col6n, near sea level, Kress & Clarkson 77-87 1 (puke), Standley 31117 (us); W of Limon Bay, Gatun ks, and Gatin Lake, Johnston 1639 (Gu); Barro Colorado Is., laboratory clearing, Croat 12422 (MO, NY, SCZ). PHENOLOGY. Flowering during wettest part of rainy season (late August prob- ably to December). HABITAT AND DISTRIBUTION. This species is found in flooded secondary growth in the Atlantic coastal lowlands of Panama (Map 7). Only a very localized population has been found west of Col6én, but this species may be scattered farther east in Panama and even into South America. 1984] KRESS, HELICONIA 493 FiGure 12. _ Heliconia sessilis. A, aang nas B, C, cincinnal bracts: B, entire (Ba, way, floral bracts removed showing flower at outer surface showing 2 ‘sepals fused to partially spread-open corolla tube. G, H, sta inode: G, abaxial view; H, position relative to style, lateral view. I, style and stigma. 494 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 T T T T 86 82 78 1 1 1 J Map 7. Costa Rica and Panama, showing restricted distribution of Heliconia sessilis RELATIONSHIPS. Heliconia sessilis is allied to members of the curtispatha and pogonantha groups with large, leafy shoots and massive inflorescences. How- ever, it most likely belongs to a separate lineage that is made up primarily of taxa found in South America. This species is characterized by its semiaquatic habitat, its attenuate leaf bases, its sessile inflorescences, and its only slightly sigmoid perianths. 14. Heliconia mariae J. D. Hooker, J. Proc. Linn. Soc., Bot. 7: 69. 1864. Type: New Granada [Colombia], Bolivar, Betami on the Sinu River (lat. 8° N), A. Anthoine s.n., 1859 (holotype, k!). PiaTE VI, B, D. Bihai mariae (J. D. Hooker) Kuntze, Rev. Gen. Pl. 2: 684. 1891. Heliconia elegans Petersen in Martius & Eichler, Fl. Brasil. 33: 12. 1890. Bihai elegans (Petersen) Kuntze, Rev. Gen. Pl. 2: 684. 1891. Type: Panama, [Canal Zone,] Paraiso, M. Wagner s.n., Dec. 1857 (holotype, M!). a Llane Griggs, Bull. Torrey Bot. Club 42: 321. 1915. Heliconia punicea (Griggs) B. Smith, Contr. Gray Herb. 124: 6. 1939. Type: Panama, Canal Zone, between ae or Gatun, elev. 10-50 m, Pittier 2290 (holotype, us!). Herb with Musa-like habit, 4-7.5 m tall. Leafy shoots in groups of 5 to 30; pseudostem red-green, glabrous, 1.9-3.8 m tall, 6-10 cm in diameter; leaves 4 to 6 per shoot; petiole green to yellow-green, often with brown spots, glabrous, 0.9-1.7 m long, 1.5-2.5 cm in diameter; blade with the base unequal, truncate, the apex obtuse to acute, the upper surface green to dark green, glabrous, with midrib light green and glabrous, the lower surface green, glabrous to slightly glaucous, midrib yellow-green with maroon midstripe and glabrous, the longest blade 1.7—2.5 m by 45-65 cm. Inflorescence pendent, long lived, to 65 cm long; peduncle red to yellow-red, densely scurfy, 20-70 cm long, 1.4-1.8 cm in diameter; rachis slightly flexuose, red to yellow, glabrous to scurfy, 1.5-1.8 cm in diameter at base; cincinnal bracts distichous, 40 to 65 per inflorescence, oriented 90-100° to axis of inflorescence, the basal bract usually fertile, the 1984] KRESS, HELICONIA 495 middle bract with apex acute, margins straight, inner surface white to pink and glabrous, outer surface pink to red (becoming yellowed with age) and glabrous to scurfy, 4.5-6.5 cm long, 7.5-9 cm wide at base, 1/w = 0.7. Floral bracts opaque, persistent, white, glabrous to puberulous abaxially, 4-5 cm long, 1.5- 2.5 cm wide at base. Flowers 20 to 30 per cincinnus; pedicel white to pink, glabrous to puberulous, 8-15 mm long; perianth white and glabrous basally, pink to red and puberulous apically, 3-4 cm long, 5-6 mm wide at base, at anthesis curved 40-70°, parabolic; free sepal slightly reflexed, fused sepals with apices reflexed; staminode white, 5-7 by 2-3 mm, apex acute to acuminate; stamens with anthers connivent inside perianth apex; pollen PCE OnmUNS Le medium size (56 x 76 um), euoblate (P/E = 0.74), heteropolar (DR/PR = 6.84) with the distal hemisphere convex (sometimes truncate), spinulose, he ae imal hemisphere planar to subconvex, psilate, the germination aperture dis- tinct, flattened and sunken; ovary white, glabrous, 6-9 by 5-6 mm. Drupes glabrous, 1-1.2 cm by 7-9 mm wide. SPECIMENS EXAMINED. Belize. TOLEDO: Columbia Forest Reserve, ca. 1-2 mi N of en- trance, Croat 24153 (mo); along Aguacate Rd. 0.5 mi from junction with San Antonio road, Proctor 35858 (Mo); Big Fall Estate, Caliente, elev. 20 m, Whitefoord 1560 (BM). DisTRICT UNKNOWN: Fairview, elev. 200 ft, Schipp 8413 (F). Guatemala. ALTA VERAPAZ: Finca Sepacuite, Cook & Griggs 787 (us); 28 mi from Tactic on road to El Estor, elev. 650 ft, Kress et al. 77-745 (DUKE). IZABAL: 6-8 km S of Modesto Mendéz, elev. 110 m, Harmon 2551] (mo); 70 mi from Flores, just below San Luis de Petén, elev. 1000 ft, Kress et al. 77-743 (DUKE); jungle bordering Quebrada Roséul, lower slopes bordering E part of Cerro Santa Cruz, NE of San Filipe, elev. 50-150 m, Steyermark 39638 (F). Honduras. ATLANTIDA: Lancetilla valley near Tela, N Coast, Pfeifer 2115 (us); vic. of Tela at sea level, Standley 53745 (F, us); on slopes near Puente Alto stop on S.F. Co R.R. E of Ceiba, elev. 900 ft, Yuncker et al. 8529 (F, GH, Ny); Puerto Sierra, near Highland Creek, Wilson 307 (ny, Us). Nicaragua. COMARCA DEL Caso: Miguel Bikou, Robbins 5863a (MO). JINOTEGA: Salto Acatula, Rio Bocay, elev. 130 m, Stevens et al. 16758 (Mo); Valle del Cua, El Cedro, elev. 700 m, Moreno 843 (Mo), 962 (MO). MATAGALPA: slope of Cerro Musun, Quebrada El Jobo (tributary of Rio Paiwas), elev. ca. 300 m, Stevens 12012 (Mo). ZELAYA: Bluefields, Cerro Maypita, 4 km ENE of Rama, elev. ca. 150 m, Proctor et al. 27357 (F, Ny); forest near bas ca. 18 km SE of La Lux, elev. 150-200 m, Bunting & Licht 579 (F, NY); 40 mi W of Rama, elev. 300 ft, Kress et al. 77-761 (DUKE); along trail from Cerro Saslaya to San José del Hormiguero, from Loma Mollejones E, halfway to San José del Hormiguero, elev. ca. 200-400 m, Stevens 7023 (DUKE); vic. of San José del Hormiguero, elev. ca. 120-150 m, Stevens 7169 (DUKE); about 8-9 km E of Rio Kukalaya on road from Puerto Cabezas to Rosita, elev. 200-300 m, Stevens 8695 (DUKE); Kururia, elev. 50 m, Pipoly 3984 (Mo); Waylawas, elev. 90-200 m, Pipoly 4197 (mo), 4343 (Mo). Costa Rica. ALAJUELA: forest near Cerro Negro and along road Kress et al. 77-794 (DUKE); road between BriBri and Bratsi, along Rio Sixaola, elev. 10- 50 m, Burger et al. 10456 (F). PUNTARENAS: Las Cruces Trop. Bot. Garden, cult., elev. 3500 ft, Kress 78-1022 (DUKE). PROVINCE UNKNOWN: near Platanillo, Toro Amarillo, 496 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 T T ] T T T 100 90 80 L l i 1 1 i Map 8. Mexico and Central America, showing distribution of Heliconia mariae. Folis 23914 (cr); elev. 31 m, Ténduz 1455] (cr). Panama. Bocas DEL Toro: vic. of Chiriqui Lagoon, Von Wedel 2905 (GH, Mo). CANAL Zone: Barro Colorado Is., Croat 6568 (mo), Croat 8689 (Mo); along K16C, vic. of Cerro Viejo, Blum 1274 (scz), Rio Providencia and ridge S of river, elev. 5-170 m, Nee & Gentry 8714 (Mo, pMa); old site of Gorgona, Maxon 6784 (us); hills near Huile around Gatin Lake, Smith et al. 3292 (F, Us); Empire to Mandingo, Piper 5437 (us); Cerro Gordo near Culebra, Standley 26000 (us); hills near Gatin, Standley 27234 (us), between Gorgona and Gattn, elev. 10-50 m, Pittier 2290 (us); Pipeline Road, elev. ca. 75 m, Kress & Cooper-Smith 76-644 (DUKE), Kress et al. 77-855 (puke), Wilbur & Weaver 11272 (pUKeE); along R.R. and Panama Canal, 6 km W of Gamboa, elev. 30-40 m, Nee 7213 (Mo). CoLon: ridge behind Garotte, D’Arcy 9323 (us); road between Ft. Sherman and Margarita, elev. 25 ft, Kress & Clarkson 80-1146 (pUKE); camino hacia San Lorenzo después de Fort Sherman, Quistgaard 3 (pMA); Rio Guanche, elev. 0-50 ft, Antonio 4821 (Mo), DARIEN: Bayano Dam Bridge, elev. 50 ft, Antonio 4517 (mo); vic. of Boca de Cupe, elev. ca. 40 m, Allen 879 (mo); Punta Guayabo Grande, elev. 0-50 m, Antonio & Hahn 4249 (mo); between Rio Jesus and Sabado, elev. ca. 100 ft, Hammel 1347 (mo); Quebrada Camachi muricate cerca casa de Bartoc, Kennedy 2831 (Mo); tributary Rio Correl6n, Sexton 21] (Mo), PANAMA: Rio Espave, Gentry 3790 (mo, pMA); 3 km S of Alcalde Diaz, elev. 410-440 m, Nee 8554 (Mo). SAN Bias: Playén Chico and vic. of San Blas, Tappakanti, Stier 167 (mo); Sasardi, elev. 20 m, Duke 10119 (Ny). PROVINCE UNKNOWN: Tabernilla, Cowell 272 (Ny); Matachin, Kuntze s.n., Aug. 1869 (Ny). PHENOLOGY. Inflorescences very long lived (up to 8 months) from first flowering to fruiting; most plants with inflorescences in various stages all year HABITAT AND DISTRIBUTION. This species is found from Belize to Panama and extends into northern South America (Map 8). It inhabits open secondary growth from sea level to 500 meters on both Atlantic and Pacific coasts, being more abundant on the wetter, Caribbean side. Plants of Heliconia mariae are soon shaded out in later successional stages and are almost never found in primary forest habitats. 1984] KRESS, HELICONIA 497 RELATIONSHIPS. Heliconia mariae is a member of the pogonantha group, but it has several characters derived within that group, including strictly distichous cincinnal bracts, parabolic flowers with red, puberulous perianths less than 4 cm in length, and pollination by territorial nonhermit hummingbirds. 15. Heliconia pogonantha Cuf. Arch. Bot. Sist. 9: 191. 1933. Type: Costa Rica, in regione Atlantica, in selva densa ad rivum prope “La Castilla— Los Negritos,” 12 km ab ore fluminis Reventazon, Cufodontis 621 (w) (type destroyed in World War II; Field Museum photograph (Neg. 30912!) remains). NeotyPe: Costa Rica, Heredia, Puerto Viejo, Finca La Selva, Daniels & Stiles 14 (F!). Herb with Musa-like habit, 4-7.5 m tall. Leafy shoots in groups of 2 to 20; pseudostem glabrous to scurfy, 1.3—4 m tall, 5-10 cm in diameter, leaves 3 to 6 per shoot; petiole light green to yellow-green, glabrous, 1.1—1.8 m long, 1.5- 3 cm in diameter; blade with the base unequal, truncate to cordate, the apex acute, the upper surface dark green, glabrous, with midrib green to yellow- green and glabrous, the lower surface light green to green, slightly glaucous, with midrib yellow-green and maroon and slightly glaucous to scurfy, the longest blade 1.2-3.3 m by 40-65 cm. Inflorescence pendent, to 1.6 m long; peduncle red to yellow, puberulous to woolly, 10-60 cm long, 1.7-2.7 cm in diameter; rachis flexuose, red to yellow, puberulous to velutinous, 1.4-2.5 cm in diameter at base; cincinnal bracts usually spirally arranged, 20 to 55 per inflorescence, oriented 100-145° to axis of inflorescence, the basal bract usually fertile, the middle bract with apex acute to acuminate, margins straight to involute at base, inner surface yellow to red and puberulous to velutinous, outer surface red (sometimes yellow near rachis) and glabrous to velutinous, 8-12.5 cm long, 9-16 cm wide at base, I/w = 0.8. Floral bracts opaque, per- sistent, white to yellow, tomentose to velutinous with golden hairs abaxially, 5-7 cm long, 1.5-2.5 cm wide at base. Flowers 10 to 30 per cincinnus; pedicel white to yellow to pink, glabrous to puberulous, 7-16 mm long; perianth yellow, glabrous basally, velutinous with golden hairs toward apex, 5-6 cm long, 9- 12 mm wide at base, at anthesis curved 45-90° and sigmoid; free sepal slightly reflexed, fused sepals with apices reflexed; staminode white, yellow at apex, 5- 9 by 2-3 mm, apex acuminate; stamens with anthers connivent inside apex of corolla tube; pollen pogonantha type, large (60-65 x 75-86 um), euoblate (to sometimes suboblate) (P/E = 0.7 1-0.84), heteropolar (DR/PR = 6.15-8.08) with the distal hemisphere convex, truncate, spinulose, the proximal hemisphere planar, psilate, the germination aperture distinct, flattened, sunken; ovary white, glabrous, 8-12 by 5-8 mm. Drupes glabrous. DistRiBuTION. Heliconia pogonantha is widespread in the Neotropics. In Cen- tral America (Map 9) the four geographically distinct varieties generally inhabit moist forested slopes and often invade disturbed open areas from sea level to 1000 m. VARIATION. The four Central American varieties of Heliconia pogonantha differ from each other in inflorescence vestiture and color and in overall size. Ad- ditional geographic variants of this species are found in South America. 498 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 65 [ —T Zz T T T T piesa ye Vv i 86 g28 78 L L iY Map 9. Nicaragua, Costa Rica, and Panama, showing distribution of Heliconia po- a: var. pogonantha (solid circles), var. holerythra (solid triangles), var. pubescens (solid squares) and var. veraguasensis (open circles). RELATIONSHIPS. This species is closely related to Heliconia ramonensis and other members of the pogonantha group. The taxa of this group differ from other heliconias with pendent inflorescences in the usually velutinous to woolly peduncles, rachises, cincinnal bracts, and perianth apices, and in the large pollen grains with flattened, sunken germination apertures. Three of the four Central American varieties of H. pogonantha have essentially glabrous inflorescences and are therefore readily distinguished from other members of the pogonantha group. NOMENCLATURE. The neotype of Heliconia pogonantha selected by Daniels and Stiles (1979) is in accordance with the protologue provided by Cufodontis. the species. The necessity for neotypification is therefore questionable. Key to the Varieties of Heliconia pogonantha A. Inflorescence entirely cee eM NES aah BS ee ANA eae ees aed 15b. var. holerythra. A. Inflorescence red and B. Peduncle, rachis, po Goan bracts velutinous to woolly. ................. B. Peduncle, rachis, and cincinnal bracts glabrous to puberulou C. Plants less than 5 m in height; lower surface of leaf blade maroon tinted; 1984] KRESS, HELICONIA 499 cincinnal bracts oriented 115-135° to axis of inflorescence at maturity. .... 15d. var. veraguasensis. C. Plants greater than 5 m in height; lower surface of leaf blade green; cincinnal bracts oriented 100-120° to axis of inflorescence at maturity. ............ et, Re ee ee eee eT CORO eres 15a. var. pogonantha. 15a. Heliconia pogonantha var. pogonantha Herb 5-7 m tall. Leafy shoots in groups of 3 to 11; pseudostem bronzy green, glabrous, 2-4 m tall, 6-8 cm in diameter; leaves 4 or 5 per shoot, petiole light green to green, 1.3-1.8 m long, 2-3 cm in diameter; longest blade 2.4-3.3 m by 41-62 cm. Inflorescence to 1.5 m long; peduncle puberulous to scurfy, 25- 55 cm long, 1.8-2.7 cm in diameter; rachis yellow, puberulous to scurfy, 1.4— 2.5 cm in diameter at base; cincinnal bracts 25 to 55 per inflorescence, oriented 100-120° to axis of inflorescence, the middle bract with margins straight to involute at base, inner surface yellow to pink and velutinous with golden hairs, outer surface red becoming yellow near rachis and puberulous to scurfy, 8—9.5 cm long, 9.5-12 cm wide at base. Floral bracts tomentose to velutinous with golden hairs abaxially, 4-5 cm long, 1.7-2.5 cm wide at base. Flowers 15 to 30 per cincinnus; pedicel pale yellow, glabrous, 9-12 mm long; perianth 5—5.8 cm long, 9-11 mm wide at base, at anthesis curved 45-70°; staminode 7-9 by 2-3 mm; ovary 1-1.1 cm by 6-8 mm SPECIMENS EXAMINED. Nicaragua. CHONTALES: ca. 4.5 km S of Hwy. 7 on road to Nueva Guinea, elev. ca. 200 m, Stevens 4949 (DUKE). Rio SAN JUAN: near Cafio Chontaleno, 20 km NE of El Castillo (Rio Indio watershed), elev. 200 m, Neil! 3425 (DUKE); in forest along Rio San Juan between San Juan del Norte (Greytown) and Delta de San Juan, elev. 0-50 m, Bunting & Licht 883 (F, Ny). ZELAYA: Bluefields, summit and slopes of Cerro San Isidro, elev. 0-65 m, Proctor et al. 27292 (F, Ny); 20 mi W of Rama, elev. 200 ft, Kress et al. 77-760 (DUKE), along road to Colonia Yolaina, signi La Esperanza, ca. 1.3 km SE of intersection with road between Nueva Guinea and Colonia Verdun, elev. 180-200 m, Stevens 6349 (Mo); along road between Rio Blanco and Rio Copalar, elev. 200-400 m, Stevens 12100 (puKE, Mo); 8 km E of bridge E of Rosita on road to Standley & Valerio 48984 (us). PUNTARENAS: Peninsula de Osa, 5 km W of Rincén de Osa, elev. 50-200 m, Burger & Gentry 9259 (F); San Vito de Java, cult. at Las Cruces Trop. Bot. Garden, elev. ca. 4000 ft, Kress 76-569 (DUKE), Kress 78-1028 (DUKE). PROv- INCE UNKNOWN: lieux humides a Lent, elev. 31 m, Ténduz 14542 (photo, Mo). PHENOLOGY. Flowering primarily during dry and early rainy seasons (January to August); some inflorescences produced nearly all year. HABITAT AND DISTRIBUTION. Heliconia pogonantha var. pogonantha is restrict- ed to the wet Atlantic lowlands below 300 m in Nicaragua and Costa Rica 500 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 north of Limén (Map 9). This variety is found in wooded late secondary growth and often develops large stands in light gaps and along forest margins. 15b. Heliconia pogonantha var. holerythra Daniels & Stiles, Brenesia 15(Supl.): 39. 1979. Type: Costa Rica, Limén, Rio Sand Box, 2 km NE of BriBri, Baker & Burger 80 (holotype, F!). Herb 4.5—7.5 m tall. Leafy shoots in groups of 2 to 18; pseudostem 1.4—2.6 m tall, 7-10 cm in diameter; leaves 3 to 6 per shoot; petiole 1.1-1.8 m long, 2-3 cm in diameter; longest blade 2—2.3 m by 40-65 cm. Inflorescence to 1.6 m long; peduncle red, scurfy, 20-60 cm long, 1.7—2.5 cm in diameter; rachis red, scurfy, 2—2.5 cm in diameter at base; cincinnal bracts 30 to 40 per inflo- rescence, oriented 1 20—140° to axis of inflorescence, the middle bract with apex acute to acuminate, often necrotic, margins straight (sometimes necrotic), inner surface puberulous, outer surface red and puberulous to scurfy, 10-12.5 cm long, 11-16 cm wide at base. Floral bracts tomentose to velutinous abaxially, 5-7 cm long, 2—2.5 cm wide at base. Flowers 15 to 30 per cincinnus; pedicel white to pink, glabrous, 1.2—1.6 cm long; perianth white to pink basally, a at apex, 5-5.7 cm long, 1-1.2 cm wide at base, at anthesis curved 7 staminode 6-8 by 3 mm; ovary white (sometimes slightly maroon), 9- 10 by 6-7 mm. Drupes 1.5-2 by 1-1.3 cm. SPECIMENS EXAMINED. Costa Rica. ALAJUELA: 38 km from Interamerican Hwy. toward Upala, Wilbur 20062 (DUKE). LIMON: 5-15 km from Siquirres on new road to Limén, elev. 50-100 m, Baker & Burger 184 (DUKE, F); Cimarrones, E of Siquirres, elev. 200 m, Maas 1134 (uv); 1-3 km N of BriBri, Rio Sixaola Drainage, elev. 20-200 m, Burger & Antonio 1098] (F); between Puerto Viejo and BriBri, elev. ca. 300 ft, Kress et al. 77- (F, MO, NY). CANAL Zone: along Transisthmian Hwy., 10 km from Coldén turnoff, elev. ca. 300 ft, Kress & Clarkson 80-1150 (DUKE); Pipeline cae Andersson & Sytsma 1233 (GB, Ss); along Rio Mendosa near Pipeline Road bridge, 8 km NW of Gamboa, elev. 95 m, Nee 7742 (us). CocLé: above La Pintada on road to Coclecito, elev. 1600-1700 ft, Kress et al. 80-1164 (DUKE). COLON: E Santa Rita Ridge lumber road, Correa & Dressler 619 (scz); Santa Rita Hills, C. E. Smith & H. M. Smith 3447 (F, us); Santa Rita Ridge, Gentry & Dwyer 9550 (mo); Santa Rita, 4 km desviacion de la Transistmica, Cuadro 1, elev. 150 m, G6mez-Pompa et al. 3222 (Mo, PMA); near Guasimo along river, Croat 9954 (mo); Rio Guanche, elev. ca. 25 m, Kress & Cooper-Smith 76-658 (DUKE); Rio Buenaventura, 3 mi from Portobelo, ca. sea level, Kress et al. 77-874 (DUKE); below Cerro Bruja, elev. 600-700 m, Andersson & Sytsma 1240 (GB, MO, S). DARIEN: trail from Punta Guayabo Grande to Rio Jaqué, elev. 50-200 m, Antonio & Hahn 4385 (mo). PANAMA: Alto de Pacora below Cerro Jefe, elev. 800-900 m, Kress et al. 76-636 (DUKE); El Llano—Carti Road, elev. 1600 ft, Kress & Hammel 82-1334 (DUKE). VERAGUAS: valley of Rio Dos Bocas on road between Alto Piedra (above Santa Fé) and Calovébora, elev. 350-400 m, Croat 27390 (Mo); valley of Rio Dos Bocas, 11 km from Escuela Agricola Alto Piedra above Santa Fé on road to Calovébora, elev. 450 m, Croat 27515 (mo); Atlantic slope, beyond Santa Fé on road to Calovébora, elev. 1200 ft, Kress et al. 80- 1199 (DUKE). PHENOLOGY. Flowering primarily during dry and early rainy seasons (January to August). HABITAT AND DISTRIBUTION. This variety is found on the Atlantic slopes and coastal plains of Costa Rica and Panama from Limén to Darién, extending 1984] KRESS, HELICONIA 501 into South America (Map 9). It is most common at elevations from sea level to 200 m but occurs to over 800 m. Variety Aolerythra prefers open disturbed habitats, often near streams or rivers. RELATIONSHIPS. Variety holerythra differs from other varieties of Heliconia pogonantha in its greater overall stature, its puberulous to scurfy, entirely red inflorescences, and its large, heavy cincinnal bracts. 15c. Heliconia pogonantha var. pubescens Daniels & Stiles, Brenesia 15(Supl.): 37. 1979. Type: Costa Rica, [Lim6én,] Turrialba—Siquirres road, 840 m elev., Daniels & Stiles 40 (holotype, F'). Herb 5-7 m tall. Leafy shoots in groups of 8 to 20; pseudostem 1.8-2.7 m tall, 6-7 cm in diameter; leaves 3 to 5 per shoot; petiole 1.3-1.6 cm long, 2- 2.5 cm in diameter; longest blade 2—2.7 m by 45-55 cm. Inflorescence to 1.5 m long; peduncle yellow to red, woolly with yellow hairs, 30-55 cm long, 1.5- 2.2 cm in diameter; rachis yellow to red, densely velutinous with yellow hairs, 1.5-2.1 cm in diameter at base; cincinnal bracts 25 to 35 per inflorescence, oriented 140-145° to axis of inflorescence, the middle bract with margins straight, inner surface pale yellow to orange and velutinous, outer surface red distally becoming yellow near rachis and velutinous, 9-10 cm long, 9-13 cm wide at base. Floral bracts 5.5 cm long, 2 cm wide at base. Flowers 15 to 25 per cincinnus; pedicel white to pink, glabrous, 1.2 cm long; perianth 6 cm long, 8-12 mm wide at base, at anthesis curved 70—90°, staminode 5-7 by 2-3 mm; ovary 1-1.2 cm by 6-7 mm. Drupes 1.5—1.7 cm by 9-11 mm. SPECIMENS EXAMINED. Costa Rica. CARTAGO/LIMON: along road between Turrialba and Siquirres, elev. 2000-2500 ft, Kress et al. 77-788 (DUKE). CARTAGO: road from Turrialba N to Pavones and Siquirres, elev. 600-700 m, Lems 5054 (ny, us); beside Rio Tepe- mechin, 4 km SE of Pejibaye, elev. 650 m, Lent 3192 (F); forest near Rio ara elev. 800 m, Lent 3737 (F); Chitaria, elev. 900 m, Maas 755 (F). LiMOn: along Hwy from Turrialba to Limén, ca. 11 mi S of Siquirres, elev. 650 m, Croat 43326 (Mo); near Guayacan, elev. 700 m, Kress 76-611] (DUKE); along Hwy. CR 233, 15 km N of Tres Equis, elev. 300 m, Lent 528 (F); Cimarrones, ca. 10 km E of Siquirres, margin of forested valley, elev. 200 m, Maas 1134 (F, Mo); along hwy. from Turrialba to Siquirres, Sheffy 35 (cr); along hwy. to Siquirres near Chitaria, Sheffy 89 (cr). PHENOLOGY. Flowering throughout year; peaking late in dry season (February to April). HABITAT AND DISTRIBUTION. Similar to the typical variety, var. pubescens 1s found in secondary growth and along forest margins. This variety is restricted to the Atlantic foothills in Costa Rica between Turrialba and Siquirres at elevations of 200-800 m (Map 9). RELATIONSHIPS. The velutinous to woolly peduncle, rachis, and cincinnal bracts distinguish var. pubescens from the other varieties of Heliconia pogonantha. 15d. Heliconia pogonantha var. veraguasensis Kress, J. Arnold Arbor. 62: 256. 1981. Type: Panama, Veraguas, 2 mi above Santa Fé beyond Agricul- tural School, elev. 850 m, Kress, Clarkson, & McDade 77-825 (holotype, DUKE!; isotypes, K!, Mo!). PLaTE VII, A, C. 502 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Herb 4—4.5 m tall. Leafy shoots in groups of 3 to 15; pseudostem 1.3-1.7 m tall, 5 cm in diameter; petiole 1-1.2 m long, 1.5-1.8 cm in diameter; leaf blade with lower surface green tinted with maroon especially along margins, longest blade 1.2-1.5 m by 40-45 cm. Inflorescence to 75 cm long; peduncle yellow, scurfy to villous, 10-30 cm long, 1.8-2 cm in diameter; rachis yellow, puberulous to villous, 1.8—-1.9 cm in diameter at base; cincinnal bracts 20 to 25 per inflorescence, oriented 115-135° to axis of inflorescence, the middle bract with margins straight, inner surface red-yellow and tomentose, outer surface red becoming yellow near rachis and glabrous to puberulous, 9—-10.5 cm long, 8-9 cm wide at base. Floral bracts pale yellow, tomentose to velutinous with golden hairs abaxially, 4—-4.8 cm long, 1.5-2.5 cm wide at base. Flowers 10 to 15 per cincinnus; pedicel white to yellow, puberulous, 7-12 mm long; perianth 5.4-5.7 cm long, 1-1.1 cm wide at base, at anthesis curved 80-90%; staminode 6-7 by 3 mm; ovary 8-9 by 5-6 mm SPECIMENS EXAMINED. Panama. VERAGUAS: vic. of Santa Fé, forested slopes of Cerro Tute, elev. 3000 ft, Allen 4333 (mo); road between Santa Fé and Rio Calovébora, 1.8 mi beyond Escuela Agricola Alto Piedra, elev. 735 m, Croat & Folsom 34255 (mo); shoulder of Cerro Tute, Folsom & Edwards 53355 (Mo); Cerro Tute, elev. 800-1350 m, 76-655 (DUKE), 76-656 (DUKE), elev. 2800-4600 ft, Kress et al. 80-1189 (DUKE); beyond Santa Fé along road to Calovébora, Pacific slopes, elev. 2200 ft, Kress et al. 80-1200 (puKE), Andersson & Sytsma 1288 (GB, s); lower montane wet forest 7 km W of Santa Fé, elev. 2900 ft, Nee 11207 (us); Caribbean slope above Rio Primero Brazo, Liesner 9 (mo); Rio Secundo Brazo, elev. 600-750 m, Knapp & Dressler 5372 (Mo), Maas & Dressler 1673 (v). PHENOLOGY. Flowering March to September; during most of year at least a few individuals reproductive. HABITAT AND DISTRIBUTION. Heliconia pogonantha var. veraguasensis is known only from middle-elevation (735-1000 m) wet forests and secondary growth in the vicinity of Cerro Tute on the Pacific slopes in Weraguas, Panama (Map 9 RELATIONSHIPS. Variety veraguasensis is distinguished by its highly reflexed cincinnal bracts, its lower leaf surfaces tinted with maroon, and its overall stature smaller than in other varieties of Heliconia pogonantha. 16. Heliconia ramonensis Daniels & Stiles, Brenesia 15(Supl.): 39. 1979. Type: Costa Rica, [Alajuela,] La Balsa, 11 mi N of San Ramon, elev. 1000 m, Daniels, Kress, & Hutchison 111 (holotype, F!; isotypes, CR, US). PLATE VII, C, D. Herb with Musa-like habit, 4-6 m tall. Leafy shoots in groups of 2 to 10; pseudostem green to yellow to brown, glabrous to villous, 1-2 m tall, 4.5-8 cm in diameter; leaves 3 to 5 per shoot; petiole rusty green to green, glabrous, 1-1.8 m long, 1.1-2 cm in diameter; blade with the base unequal, obtuse to cordate, the apex acute to acuminate, the upper surface dark green, glabrous, 1984] KRESS, HELICONIA 503 with midrib light green to yellow-green and glabrous, the lower surface green to maroon, glabrous, with midrib green to maroon and glabrous, the longest blade 1-2 m by 30-60 cm. Inflorescence pendent, to 90 cm long; peduncle pink to rose-red to red, glabrous to woolly with golden or rusty orange hairs, 20-60 cm long, 1.3-2 cm in diameter; rachis flexuose, red to pink to yellow, glabrous to woolly, 1-2.2 cm in diameter at base; cincinnal bracts distichous to subspirally arranged, 20 to 40 per inflorescence, oriented 120-140° to axis of inflorescence, the basal bract usually fertile, the middle bract with apex acute to acuminate, margins straight to involute at base, inner surface pink to red to yellow and velutinous to villous with golden or rusty hairs, outer surface red to rose-red to pink and glabrous to woolly, 7-12 cm long, 7-10 cm wide at base, I/w = 1.1. Floral bracts opaque, persistent, white to pink, glabrous to villous abaxially, 4-5 cm long, 1.5—3 cm wide at base. Flowers 10 to 30 per cincinnus; pedicel white to pink, glabrous, 0.7-3 cm long; perianth white to light pink and glabrous basally, yellow and velutinous toward apex, 4.5-5.6 cm long, 1-1.2 cm wide at base, at anthesis curved 60-90° and sigmoid; free sepal reflexed, fused sepals with apices reflexed; staminode white (sometimes with yellow apex), 6-10 by 2-4 mm, apex acuminate to cuspidate; stamens with anthers connivent inside corolla apex; pollen pogonantha type, large (60- 63 x 78-88 wm), euoblate (P/E = 0.68-0.81), heteropolar (DR/PR = 5.79-8.7) with the distal hemisphere convex, truncate, spinulose, the proximal hemi- sphere planar, psilate, the germination aperture distinct, flattened, sunken; ovary white to pale blue to lavender, glabrous, 7-10 by 5-7 mm. Drupes glabrous, 1.3-1.5 cm by 8-10 mm; pyrenes 9 by 5 mm. HABITAT AND DISTRIBUTION. Heliconia ramonensis occurs in middle-elevation (600-1200 m) forests of Costa Rica and Panama (Map 10). Plants inhabit moist, forested slopes and often invade disturbed open areas. VARIATION. The four varieties of Heliconia ramonensis are distinguished from each other by the distribution of the woolly vestiture on the peduncle, rachis, and cincinnal bracts, and by the color of the hairs on the floral bracts and perianth apices. RELATIONSHIPS. Heliconia ramonensis belongs to the pogonantha group; it differs from other members of that group in the rusty orange hairs on various parts of its inflorescence and peduncle and in its bright pink to red cincinnal bracts. Key to the Varieties of Heliconia ramonensis A. Peduncle, rachis, and cincinnal bracts all essentially glabrous. .... 16c. var. glabra. A. Peduncle, rachis, or cincinnal bracts villous to woolly. B. Peduncle, rachis, and cincinnal bracts all woolly. ........ 16d. var. /anuginosa. B. Peduncle and sometimes rachis woolly, cincinnal bracts glabrous to villous. C. os on floral bracts and perianth pale yellow to buff, cincinnal bracts br ryan inioid grat gsa Ned iad acpeect ns ear ees apes a etenie eatin de a. var. ramonensis. Cc; Vestine on floral bracts and perianth bright yellow; cincinnal bracts rose- red to pink. 0.0.0.0... eee 16b. var. xanthotricha. 504 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 T T T T 86 1 Map 10. Costa Rica and Panama, showing allopatric distribution of Heliconia ra- nensis var. ramonensis (solid circles), var. xanthotricha (solid triangles), var. glabra (solid square), and var. /anuginosa (open circles). 16a. Heliconia ramonensis var. ramonensis Herb to 5 m tall. Pseudostem 1.5-2 m tall, 4.5-6 cm in diameter; leaves with the petiole rusty green, 1.4-1.8 m long, 1.7—1.9 cm in diameter, the blade with base obtuse to truncate, the longest blade 1.6-2 m by 45-55 cm. Inflo- rescence to 85 cm long; peduncle red, woolly with golden hairs, 40-60 cm long, 1.8-2 cm in diameter; rachis red, villous to woolly with golden hairs, 1.5-2 cm in diameter at base; cincinnal bracts 20 to 30 per inflorescence, the middle bract with inner surface villous with golden hairs, outer surface red to pink- red and puberulous to scurfy, 9-10 cm long, 8-10 cm wide at base. Floral bracts pink, glabrous to villous with golden hairs abaxially, 4-5 cm long, 1.5- 3 cm wide at base. Flowers 20 to 30 per cincinnus; pedicel 1.2-3 mm long; perianth glabrous basally, villous with golden hairs toward apex, 4.5-5.5 cm long, 1-1.2 cm wide at base; staminode white, yellow at apex, 7-10 by 3-4 mm; ovary pale lavender, 7-10 by 6-7 mm. Drupes not seen. SPECIMENS EXAMINED. Costa Rica. ALAJUELA: Cataratas de San Ramon, Brenes 18370 (cr); 15 mi from San Ramon near La Balsa, elev. 1000 m, Kress et al. 76-604 (DUKE), 77-764 (DUKE); entre Balsa de San Ramon y Rio Cataratas, ca. 12 km N of La Balsa, elev. 550-1100 m, Utley & Utley 3745 (DUKE). PHENOLOGY. Flowering primarily during early rainy season (July to September), some inflorescences produced throughout year. DISTRIBUTION. Heliconia ramonensis var. ramonensis is endemic to Costa Rica in the foothills (ca. 1000 m) above San Ramén (Map 10). 16b. Heliconia ramonensis var. xanthotricha Kress, J. Arnold Arbor. 62: 258. 1981. Type: Panama, Coclé, 6 mi beyond El Valle de Antén behind Cerro Gaital, elev. 900 m, Kress, Clarkson, & McDade 77-840 (holotype, DUKE!; isotypes, K!, Mo!). 1984] KRESS, HELICONIA 505 Heliconia vellerigera auct, non Poeppig: Woodson & Schery, Ann. Missouri Bot. Gard. 5, in part (Woodson & Schery included H. magnifica and H. ramonensis in H vellerigera). Herb 4—5 m tall. Leafy shoots in groups of 3 to 10; pseudostem orange to brown, 1.3—2 m tall, 5-8 cm in diameter; leaves with the petiole green, 1-1.2 m long, 1.5—2 cm in diameter, the blade with base truncate to cordate, lower surface green tinted with maroon especially along margins, the longest blade 1.3-1.8 m by 40-60 cm wide. Inflorescence to 90 cm long; peduncle red, woolly with golden-yellow hairs, 30-50 cm long, 1.6-—2 cm in diameter; rachis red- orange, velutinous to villous with yellow hairs, 1.5—2.2 cm in diameter at base; cincinnal bracts 20 to 40 per inflorescence, oriented 135-140° to axis of inflo- rescence, the middle bract with inner surface pink and velutinous to villous, outer surface rose-red, 9-12 cm long, 7-10 cm wide at base. Floral bracts white to pink, velutinous to villous with bright yellow hairs abaxially. Flowers 10 to 15 per cincinnus; pedicel 7-13 mm long; perianth glabrous basally, tomentose to velutinous with bright yellow hairs toward apex, especially along sepal mar- gins, 5-6 cm long, 1-1.2 cm wide at base, at anthesis curved 90°; staminode 7-8 by 2-3 mm; ovary bluish white, 8-10 by 5-6 mm. Drupes 1.3-1.5 cm by 9-10 mm. SPECIMENS EXAMINED. Panama. Coc_é: base of three peaks beyond El Valle de Antén, 1000 m, Allen 2707 (Mo), Allen 2867 (us); vic. Finca Tomas Arias, elev. 600 m, Allen 3629 (8) foot of Cerro Pilén, elev. 2000 ft, Porter et al. 4423 (scz, uc), Croat 13471 (mo); La Mesa, 2.8-3.4 mi NW of El Valle de Antén, elev. 850-900 m, Luteyn 4064 (DUKE), Croat 25311 (mo), elev. 900 m, Croat 14374 (Mo, scz), 22952 (Mo), ean 1292 (mo), elev. 900 m, Sullivan 541 (mo), D’Arcy & Sytsma 3 ae ca. 9k El Valle Market beyond Sr. Furlong’s Finca, Kennedy et al. 3223 (DUKE, an ar erro Gaital, N of El Valle, elev. 760 m, Reveal & Balogh 4971 ao “ley 2200 a ae et al. 80-1161 (DUKE); near Cerro Turega, elev. 650-700 m, Woodson & Schery 205 (mo); erro Caracoral, elev. 2700-3200 ft, Sytsma 3764 (Mo), Bee ao. 4053 (MO), penne & Sytsma 1249 (s). PHENOLOGY. Flowering mainly during rainy season (May to September); some inflorescences produced throughout year. DIsTRIBUTION. In Central America Heliconia ramonensis var. xanthotricha 1s found at middle elevations (650-1000 m) in Coclé, Panama, in the region around El Valle de Anton (Map 10). RELATIONSHIPS. Variety xanthotricha differs from vars. ramonensis and lan- uginosa in the bright yellow vestiture on its floral bracts and perianth apices (also found in var. glabra), and in the contrasting colors of its red-orange rachis and rose-red cincinnal bracts. l6c. Heliconia ramonensis var. glabra Kress, var. nov. A var. ramonensi inflorescentiis glabris et floribus pubescentiis luteis differt. Type: Panama, Coclé, beyond sawmill above El Copé, elev. 2400 ft, Kress & Hammel 83-1589 (holotype, DUKE!; isotype, Mo!). 506 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Herb 4-5 m tall. Leafy shoots in groups of 5 to 10; pseudostem green and brown, glabrous, 1-1.5 m tall, 5-7 cm in diameter; leaves with the petiole green, 1-1.2 m long, 1.1—1.5 cm in diameter, the blade with base cordate, apex acute, lower surface tinted maroon with maroon lateral venation and green midrib, the longest blade 1.1-1.3 m by 30-40 cm. Inflorescence to 50 cm long; peduncle rose-red, glabrous, 20-30 cm long, 1.3-1.5 cm in diameter; rachis rose-red, glabrous to scurfy, |-1.3 cm in diameter at base; cincinnal bracts 20 to 30 per inflorescence, oriented 135° to axis of inflorescence, the middle bract with inner surface pink and velutinous with yellow hairs, outer surface rose- red to pink and glabrous, 7-8 cm long, 7-9 cm wide at base. Floral bracts white, villous with bright yellow hairs abaxially, 5 cm long, 2 cm wide at base. Flowers 25 per cincinnus; pedicel pink, | cm long; perianth with bright yellow hairs toward apex, 5.6 cm long, 1.2 cm wide at base, at anthesis curved 85°; staminode 6 by 2 mm; ovary white, 8 by 6 mm. Drupes not seen. SPECIMENS EXAMINED. Panama. Coc .é: area surrounding Rivera sawmill, 7 km N of El Copé, continental divide, elev. 750-860 m, Folsom & Collins 6430 (DUKE, MO). PHENOLOGY. Apparently flowering mainly during rainy season (July to No- vember). DISTRIBUTION. Although quite common at the type locality (pers. obs.), var. glabra has been collected only a few times and appears to be restricted to the mountains (750-850 m) above El Copé in Coclé, Panama (Map 10). RELATIONSHIPS. This variety is quite different from the other varieties of Hel- iconia ramonensis in its entirely glabrous peduncle, rachis, and cincinnal bracts. The bright yellow hairs of the floral bracts and perianths are also present in var. xanthotricha. 16d. Heliconia ramonensis var. lanuginosa Kress, var. nov. PLATE VII, B, D. A var. ramonensi inflorescentiis omnibus lanuginosis et pubescentia ferru- ginea differt. Tyre: Panama, Chiriqui, La Fortuna Dam site above Gualaca, ee 3600 ft, Kress & Hammel 83-1602 (holotype, DUKE!; isotypes, K!, Mo!, PMA!). Herb 5-6 m tall. Leafy shoots in groups of 2 to 5; pseudostem yellow to brown, glabrous to villous, 2 m tall, 5-6 cm in diameter; leaves with the petiole green and maroon, |-1.2 m long, 1.6-1.8 cm in diameter, the blade with base truncate to cordate, lower surface green (sometimes with maroon borders), the longest blade 1.6-1.8 m by 37-44 cm. Inflorescence to 75 cm long; peduncle pink, woolly with rusty orange hairs, 20-50 cm long, 1.5-1.7 cm in diameter; rachis pink to pale yellow, woolly with rusty orange hairs, 1.4 cm in diameter at base; cincinnal bracts 20 to 25 per inflorescence, oriented 120-135° to axis of inflorescence, the middle bract with inner surface pale yellow and villous to velutinous with rusty orange hairs, outer surface pink to rose-red and villous to woolly with rusty orange hairs, 10-11 cm long, 7-9 cm wide at base. Floral bracts white, velutinous with rusty orange hairs abaxially, 4-4.5 cm long, 1.9- 2 cm wide at base. Flowers 20 per cincinnus; pedicel white, 5-8 mm long; 1984] KRESS, HELICONIA 507 perianth light pink basally, velutinous with rusty orange hairs apically, 5—5.5 cm long, 1.2 cm wide at base, at anthesis curved 90°; staminode white with yellow apex, 8-9 by 2 mm, apex acuminate; ovary white, 6-10 by 5 mm. SPECIMENS EXAMINED. Panama. CHIRIQUI: along road between Gualaca and Fortuna Dam site, 7.9 mi NW of Los Planes de Hornito, elev. 1300 m, Antonio 4130 (mo); N del sitio de presa de Fortuna, elev. 1000-1200 m, Correa et al. 2317 (Mo), 2670 (Mo); Fortuna Dam site, elev. 1040 m, Hammel 1989 (mo), elev. 1000-1200 m, Kress et al. 82-1350 (DUKE), Kress & Hammel 83-1600 (DUKE), elev. 1200-1400 m, Folsom et al. 5493 (mo). Bocas DEL Toro: Atlantic slopes beyond La Fortuna Dam site, elev. 1000 m, Kress & Hammel 83-1604 (DUKE). PHENOLOGY. Flowering mainly during rainy season (May to September). DisTRIBUTION. Variety /anuginosa has been collected in the mountains (1000- 1300 m) surrounding the Fortuna Hydroelectric Dam project in Chiriqui and neighboring Bocas del Toro, Panama (Map 10). RELATIONSHIPS. The inflorescences of this variety are conspicuously different from those of the other varieties in being entirely covered with a woolly in- dumentum of rusty orange hairs. In vars. ramonensis and xanthotricha this rusty vestiture is restricted mainly to the peduncle and rachis. The rusty orange hairs on the floral bracts and perianth of var. /anuginosa differ from the yellow hairs of the other varieties. 17. Heliconia danielsiana Kress, sp. nov. FiGureE 13; PLate VIII, A, D. Heliconia vellerigera auct. non Poeppig: Daniels & Stiles, Brenesia 15(Supl.): 45. 1979. Planta herbacea, 4.5-8 m alta; inflorescentia pendens, rubra, pubescentia lanata aurantiaca usque ad bubalinam; cincinnorum bracteae distichae vel subspirales; perianthium basi album apice lutescens, apice pubescentia cin- namomea usque ad bubalinum; ovarium album glabrum; fructus caeruleus glaber Type: Costa Rica, San José, 15 km above Dominical on road to San Isidro, elev. 2800 ft, Kress & Clarkson 79-1097 (holotype, DUKE!; isotypes, cR!, F!). Herb with Musa-like habit, 4.5-8 m tall. Leafy shoots in groups of 5 to 15; pseudostem brown, scurfy to villous, 1.8-3 m tall, 5-8 cm in diameter; leaves to 4 per shoot; petiole green to yellow, glabrous, 1-1.8 m long, 2—-2.5 cm in diameter; blade with the base unequal, truncate to cordate, the apex acute, the upper surface green to dark green, glabrous, with midrib green to yellow and glabrous, the lower surface green, slightly glaucous, midrib green to yellow with maroon midstripe and glabrous, the longest blade 2-2.8 m by 45-57 cm. Inflorescence pendent, to 1 m long; peduncle orange to red, woolly with orange (when fresh) to buff (when dried) hairs, 10-40 cm long, 1.6-2 cm in diameter; rachis flexuose, orange to red, woolly, 1.6—2 cm in diameter at base; cincinnal bracts distichous to subspirally arranged, 20 to 30 per inflorescence, oriented 130-150° to axis of inflorescence, the basal bract usually fertile, the middle bract with apex acute to acuminate, margins straight to involute, inner surface pink and velutinous, outer surface orange-red and woolly with orange (when 508 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Ficure 13. Heliconia danielsiana. A, inflorescence. B, C, cincinnal bracts: B, entire ow G, staminode: F, abaxial view; G, position relative to style, lateral view. H, style and fresh) to buff (when dried) hairs, 9.5—11 cm long, 8-10 cm wide at base, l/w = 1.1. Floral bracts opaque, persistent, white to pink, velutinous to villous with ares hairs abaxially, 4-5 cm long, 2-2.5 cm wide at base. Flowers 15 to 20 per cincinnus; edie white to a glabrous to puberulous, 9-12 mm long; perianth white and glabrous basally, yellow and velutinous with cinnamon to 1984] KRESS, HELICONIA 509 T T U T T 6 82 78 4 | i 1 | Map 11. Costa Rica and Panama, showing allopatric distribution of Heliconia dan- ielsiana (triangles), H. magnifica (circles), and H. xanthovillosa (squares). buff hairs apically, 5.1-5.7 cm long, 1-1.2 cm wide at base, at anthesis curved 85-90° and sigmoid; free sepal slightly reflexed, fused sepals with apices slightly reflexed; staminode white with yellow apex, 5-7 by 2-3 mm, apex acuminate; stamens with anthers connivent inside perianth apex; pollen pogonantha type, large (60 x 86 um), euoblate (P/E = 0.71), heteropolar (DR/PR = 7.9) with the distal hemisphere convex, truncate, spinulose, the proximal hemisphere planar, psilate, the germination pore distinct, flattened, sunken; ovary white, glabrous, 8-10 by 5-6 mm. Drupes glabrous. SPECIMENS EXAMINED. Costa Rica. PUNTARENAS: Golfo Dulce area, vic. Esquinas Exper- iment Station, sea level, Allen 5235 See entre Los Rios Piedras Blancas y Esquinas, Osa, elev. 75 m, Jiménez 2248 (F, GH, NY); Esquinas Forest Reserve between Palmar and Rio Claro, elev. ca. 100 ft, Kress et al. 76-589 (DUKE), Kress 78-1046 (DUKE); forest 30 km SE of Palmar Sur on Pan Am Hwy., Lent 171 (cr); forest of Rio Convento, 36 km S of San Isidro del General, elev. 500 m, Molina et al. 17980 (Ny); 8 mi from Villa Neily on road to San Vito, elev. 3500 ft, Kress et al. 77-82] (DUKE); San Vito de Java, cult. at Las Cruces Trop. Bot. Garden, elev. ca. 4000 ft, Kress 76-575 (DUKE), 78-1031 (DUKE); Golfito, along trail to TV tower, steep forested slopes, elev. 100-300 m, Burger & Matta 4772 (cr), 4773 (CR, F, GH, US); Peninsula de Osa, Corcovado Natl. Park, elev. 20-400 m, Liesner 3177 (Mo); Peninsula de Osa, Camino de Altura, 2-5 mi W of Rincén de Osa, elev. 400-1000 ft, Raven 21494 (cr, F). SAN José: El General Valley, vic. of San Isidro del General, elev. 500 m, Molina et al. 17980 (F); between San Isidro and Playa Dominical, Sheffy 62 (cr). PHENOLOGY. Flowering throughout year, with slight peak in early rainy season (June to August). HABITAT AND DISTRIBUTION. Heliconia danielsiana is restricted to Costa Rica, where it occurs only on the Pacific slopes from Dominical to the Golfo Dulce area (Map 11). This species inhabits open disturbed areas from sea level to 1200 m. 510 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 RELATIONSHIPS. This species, a member of the pogonantha group, has extremely woolly inflorescences. It is distinguished by its orange-red cincinnal bracts and by the orange to buff vestiture densely covering the entire inflorescence. NOMENCLATURE. Daniels and Stiles (1979) applied the name Heliconia velle- rigera Poeppig to the Costa Rican species characterized by an extremely woolly vestiture covering the entire pendent inflorescence. Indeed, Poeppig had de- scribed such a plant from Peru and deposited a type at Vienna (w). The type specimen is minimal, consisting only of several cincinnal bracts, and does not have any flowers or leaves. Other collections seen from Peru (e.g., Ferreyra 1198, us; Berlin 383, F; Plowman & Schunke 11471, puke, F) and Colombia (Foster & Foster 2270, GH) with similar woolly inflorescences have glabrous flowers that differ from the velutinous flowers of plants found in Costa Rica. The large geographic distance between the type locality of H. vellerigera and Costa Rica and the difference in the vestiture of the flowers leave no doubt that the plants found in Costa Rica are specifically distinct from Poeppig’s taxon. The specific epithet proposed here honors Dr. Gilbert S. Daniels, who has contributed much to our understanding of the Heliconia taxa of Costa Rica, and who first showed me the new species in its native habitat. 18. Heliconia magnifica Kress, J. Arnold Arbor. 62: 246. 1981. Type: Panama, Panama, Cerro Campana, beyond Su Lin Motel, elev. 1000 m, Kress, Clarkson, & McDade 77-850 (holotype, DUKE!; isotypes, K!, Mo!). Ficure 14; PLate VIII, B, E. Heliconia vellerigera auct. non Poeppig: Woodson & Schery, Ann. Missouri Bot. Gard. 32: 54. 1945, in part (Woodson & Schery included H. magnifica and H. ramonensis var. xanthotricha in H. vellerigera). Herb with Musa-like habit, 4-6 m tall. Leafy shoots solitary or in groups of 2 to 5; pseudostem rusty brown, scurfy to villous, 1.1-2.3 m tall, 3-7 cm in diameter; leaves 3 or 4 per shoot; longest petiole light green and maroon, glabrous, 1.1-1.8 m long, 1-1.9 cm in diameter; blades splitting into narrow lateral segments with age, the base unequal, truncate to cordate, the apex acute, the upper surface dark green, glabrous, with midrib yellow-green and glabrous, the lower surface green with maroon margins, glabrous, with midrib yellow- green and glabrous, the longest blade 1.1-1.9 m by 36-55 cm. Inflorescence pendent, to 94 cm long; peduncle dark red to burgundy, woolly with golden to burgundy hairs, 30-50 cm long, 1-1.7 cm in diameter; rachis flexuose, twisted, red to burgundy, woolly, 1—1.6 cm in diameter at base; cincinnal bracts spirally arranged, up to 35 per inflorescence, oriented 130—150° to axis of inflorescence, the basal bract usually fertile, the middle bract with apex acuminate, margins straight becoming involute at base, inner surface white to red and pubescent to velutinous, outer surface red to burgundy and villous to woolly with bur- gundy hairs, 8-10 cm long, 7.5-9.5 cm wide at base, I/w = 1.1. Floral bracts opaque, persistent, pink, velutinous abaxially, 3.5-5 cm long, 1-1.5 cm wide at base. Flowers 10 to 15 per cincinnus; pedicel white, pink near ovary, glabrous, 1-1.3 cm long; perianth white and glabrous basally, yellow and velutinous with 511 KRESS, HELICONIA 1984] FiGure 14. Heliconia magnifica. A, inflorescence. B, C, cincinnal bracts: B, entire a, flower protruding at anthesis); C, cut-away, floral bracts removed showing flower perianth, outer surface showing 2 sepals fused to partially spread-open corolla tube. G, H, staminode: G, abaxial view; H, position relative to style, lateral view. I, style and stigma. J, mature leaf, blade split into lateral segments. 512 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 golden hairs toward apex, 5.5-6 cm long, |-1.2 cm wide at base, at anthesis curved 80—-90° and sigmoid; free sepal reflexed, fused sepals with apices slightly reflexed; staminode white, yellow at apex, 5-6 by 2-3 mm, apex acuminate; stamens with anthers connivent inside apex of corolla tube; pollen pogonantha type, large (65 x 82 um), suboblate (P/E = 0.78), heteropolar (DR/PR = 6.07) with the distal hemisphere convex, truncate, spinulose, the proximal hemi- sphere planar, psilate, the germination aperture distinct, flattened, sunken; ovary white, glabrous, 1-1.1 cm by 5-6 mm. Drupes glabrous, 1.2-1.3 cm by 8-10 mm; pyrenes 9 by 6 mm. SPECIMENS EXAMINED. Panama. PANAMA: Cerro Campana, elev. 600 m, Antonio 1250 (Mo), 1725 (Mo), 4855 (mo), elev. 700-800 m, Andersson 1317 (Gs, s), elev. 850 m Busey 847 (Mo), Liesner 627 (F, MO, NY), elev. 875 m, Croat 25239 (Mo), elev. 2600- 2800 ft, Lewis et al. 1903 (Mo), elev. 800-1000 m, Gentry 4955 (F, Mo, scz), elev. 1000 m, Allen 2425 (Mo, s), near summit, elev. 1000 m, Kress & Cooper-Smith 76-648 (DUKE), elev. 2800 ft, Sytsma 1233 (mo), elev. 2700-3200 ft, Hammel 3783 (Mo), elev. 3300 ft, Kress 80-1246 (DUKE), ¥% of way to summit from Pan American Hwy., Dwyer et al. 4850 (Mo), cloud forest, Luteyn & Kennedy 1814 (DUKE), without further location, C. E. Smith & H. M. Smith 3399 (us). PHENOLOGY. Flowering mainly March to September; some mature inflores- cences all year. HABITAT AND DISTRIBUTION. This species is found at middle elevations in the understory of cloud forest. It is endemic to Cerro Campana in Panama at elevations above 600 m (Map 11) RELATIONSHIPS. Heliconia magnifica 1s closely related to other species of the pogonantha group with large pendent woolly inflorescences, including H. dan- ielsiana, H. ramonensis, and H. xanthovillosa, found in Central America. It is distinguished from these species by the dark red to burgundy color of its inflorescence and the red to golden woolly pubescence covering its peduncle, rachis, and cincinnal bracts. 19. Heliconia xanthovillosa Kress, J. Arnold Arbor. 62: 253. [21 Aug.] 1981. : Panama, Panama, El Llano—Carti Road, 12 km from intersection with Pan American Hwy., elev. 500 m, Kress & Knapp 80-1233 (ho- lotype, DUKE!; isotypes, F!, GH!, K!, MOo!, PMAa!). FiGurE 15; PLATE VIII, C, F. Heliconia dresslerana Abalo & Antonio, Baileya 21: 139. [17 Sept.] 1981. Type: Pan- ama, Panama, El Llano-Carti Road, 15 km N of Panamerican Hwy. at El Llano, Antonio 5258 (holotype, Mo!; isotypes, PMA, US). Herb with Musa-like habit, 3.5—4.5(-7) m tall. Leafy shoots in groups of 2 to 11; pseudostem brown, scurfy to puberulous, 1.6—1.9(-—3.5) m tall, 6-7 cm in diameter; leaves 4 per shoot; petiole dull green, glabrous, 1.1-1.2 m long, 1.9-2(-2.5) cm in diameter; blade with the base unequal, obtuse to truncate, the apex acute, the upper surface green to dark green, glabrous, with midrib light green and glabrous, the lower surface green, glabrous, with midrib yellow- green and glabrous, the longest blade 1.9-2.3 m by 49-55 cm. Inflorescence 1984] KRESS, HELICONIA le scence. B, C, cincinnal bracts: B, entire floral bracts removed showing flower Figure 15. Heliconia xanthovillosa. A, inflore (Ba, flower protruding at anthesis); C, cut-away, included in apex of corolla tube). D, E, : D, perianth, outer surface showing 2 sepals fused to partially spread-open corolla tube. G, H, staminode: G, abaxial view; H, position relative to style, lateral view. I, style and stigma. 514 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 pendent, to 86(-105) cm long; peduncle bright yellow-green, woolly with yellow hairs, 26-68 cm long, 1.5-2.2 cm in diameter; rachis flexuose, bright yellow, woolly with yellow hairs, 1.5-1.9 cm in diameter at base; cincinnal bracts distichous to subspirally arranged, up to 35 per inflorescence, oriented 135- 145° to axis of inflorescence, the basal bract sterile or fertile, the middle bract with apex acuminate to acute, margins straight to involute at base, inner surface pale yellow-green (sometimes pink) and villous, outer surface bright yellow to green-yellow (sometimes red distally) and woolly with yellow hairs, 8-10 cm long, 7-8 cm wide at base, l/w = 1.2. Floral bracts opaque, persistent through fruiting, pale yellow, velutinous to villous abaxially, 3.7-4.8 cm long, 1.6-1.7 cm wide at base. Flowers 12 to 20 per cincinnus; pedicel white, glabrous (to hirsute with yellow hairs), 1 cm long; perianth white and glabrous basally, yellow and velutinous to villous with bright yellow hairs distally, 5.1-5.5 cm long, 9-11 mm wide at base, at anthesis curved 85—-90° and sigmoid; free sepal reflexed, fused sepals with apices slightly reflexed; staminode white, 5(-8) by 3(-5) mm, apex acuminate; stamens with anthers connivent inside apex of corolla tube; pollen pogonantha type, large (65 x 81 um), suboblate (P/E = 0.79), heteropolar (DR/PR = 6.63), with the distal hemisphere convex, trun- cate, spinulose, the proximal hemisphere planar, psilate, the germination aperture distinct, flattened, sunken; ovary white, glabrous, 7-8 by 5-6 mm. Drupes glabrous, 1.1-1.2 cm by 8 mm; pyrenes 8-9 by 5-6 mm. SPECIMENS EXAMINED. Panama. PANAMA: El Llano—Carti Road, 8 mi from Pan American Hwy., elev. 1200 ft, Antonio 1699 (mo), elev. 300-500 m, Folsom & Maas 5214 (MO), Antonio 1266 (Mo), 1277 (Mo), Folsom 3507 (Mo), Maas et al. 2810 (GB), Andersson & Sytsma 1262 (Gp, s), Knapp et al. 4747 (mo), Kress & Hammel 82-1335 (puKe), 83- 1564 (DUKE), 83- 1565 (DUKE), Kress & Knapp 80-1235 (DUKE). PHENOLOGY. Flowering during rainy season (May to October); fruiting probably well into November. HABITAT AND DISTRIBUTION. This species is found at middle elevations (300- 500 m) in open areas in the forest, and now especially in areas disturbed by man. It has been collected only along the continental divide separating the drier Pacific slopes from the wetter Atlantic ones in eastern Panama above the Rio Carti (Map 11). VARIATION. Several individuals having large, stout inflorescences with yellow and red cincinnal bracts (Antonio 1277; Kress & Knapp 83-1235: Kress & Hammel 82-1335, 83-1564) have been collected on El Llano-Carti Road. Although differing in bract color, the inflorescences are covered with the same yellow vestiture as typical Heliconia xanthovillosa and are thus considered a color form of that species. RELATIONSHIPS. Heliconia xanthovillosa is closely related to species of the po- gonantha group, especially H. danielsiana, H. ramonensis, and H. magnifica, which differ from each other essentially in cincinnal-bract and indumentum color. Heliconia xanthovillosa has bright yellow inflorescences that are entirely covered with long, yellow hairs. 1984] KRESS, HELICONIA 515 INSUFFICIENTLY KNOWN TAXON In the examination of numerous herbarium specimens, several collections from Panama (one from the Burica Peninsula in Chiriqui and two from Darién) have proved quite difficult to assign to any of the taxa of Heliconia with pendent inflorescences included here. These specimens have a characteristic inflores- cence with a strongly flexuose rachis and nearly overlapping cincinnal bracts. The cincinnal bracts are distinctive in having apparently necrotic apices that are either black or green (due to algal growth?). Dodson and Gentry (1978) have described a species from Ecuador, H. nigripraefixa, that has the same feature. I do not think that these specimens should be placed in that taxon until further comparative studies can be made on living specimens in the field. SPECIMENS EXAMINED. Panama. CHiRIQUi: Peninsula de Burica, 20 km W of Puerto Armuelles, elev. 400-600 m, Busey 551 (Mo). Darién: headwater of Rio Tuquesa, ca. km from continental divide, vicinity gold-mining camp of Tyler Kittredge, Croat 27140 (mo), 2714] (mo) NATURAL HyBRIDS Natural hybrids are rare in Heliconia. As discussed earlier, attempts to ar- tificially hybridize a wide range of heliconias have proved unsuccessful. Strong physiological stigma/style barriers to foreign pollen penetration are present in most species (Kress, 1981b, 1983b). Although a limited number of hybrids have been found involving species with pendent inflorescences (TABLE 7), only two—H. curtispatha x H. pogonantha var. holerythra and H. mariae x H. pogonantha var. holerythra—are known to occur in more than one locality. Detailed morphological descriptions and ae of the three parent taxa and their two hybrid offspring are given in TABLE 8. The hybrid nature of the plants listed in TapLe 7 (excluding those cited by TaBLe 7. Natural hybrids of Heliconia involving species with pendent inflorescences. Hysrip VOUCHER NUMBER* OR COMBINATION COUNTRY LITERATURE CITATION H. curtispatha x Panama Kress 80-1144, 80-1156 H. pogonantha H. mariae < Costa Rica Kress as ie 76-659, 77-795, 77-875 H. pogonantha William H. nutans =< Panama Kress & re 79-1089c¢ H. lankesteri H. secunda x Costa Rica Kress 77-777 H. clinophila H. stilesii x Costa Rica Daniels & Stiles (1979) H. danielsiana H. tortuosa x Costa Rica Daniels & Stiles (1979) H. nutans *For collection localities and herbaria, see “Specimens Examined” under each hybrid in text. TasLe 8. Morphological comparison of Heliconia curtispatha, H. pogonantha var. holerythra, H. mariae, and their putative hybrids. H. POGONANTHA CHARACTER H. CURTISPATHA “HYBRID” VAR. HOLERYTHRA “HYBRID” H. MARIAE Height (m) 6-7 7-8 4.5-7.5 6 4-7.5 Petiole Length (m 0.9-1.5 1.7-2 1.1-1.8 1.2-1.4 0.9-1.7 Diameter (cm) 1.52.3 2.2 2-3 2.2-2.6 1.5-2.5 Leafblade Lower surface Glaucous Glaucous Slightly glaucous Glabrous Glabrous Length (m) 1.9-2.3 2.8-3 2-3. 2.3-2.7 1.7-2.5 Width (cm) 35-55 50-60 40-65 50-60 45-65 Peduncle vestiture Absent to scurfy Scurfy Scurfy Scurfy Densely scurfy Inflorescence length (exclud- 1.6 1.5 1.6 1 0.65 ing peduncle) (m) Rachis Vestiture Puberulous to Puberulous to Scurfy Scurfy Absent to scurfy scu scurfy Diameter (cm) 1.2-1.8 2-2.5 1.8-2.2 1.5-1.8 Cincinnal bracts Arrangement Distichous to spiral Spiral Spiral Distichous to spiral Distichous Orientation (to inflores- 35-50° 50-60° 40-60° 40-50° 80-90° cence axis Inner surface color Yellow-orange to Yellow to red Yellow-red White to yellow to White to pink pin ink Inner surface vestiture Tomentose to velu- Puberulous to pu- Puberulous Absent to puberu- Absent inous bescent lous Outer surface color Red to yellow- Rose-red Red Red Pink to red Outer surface vestiture Puberulous Absent Puberulous Puberulous to scur- Absent to scurfy WOLAYOUUV GATIONAV AHL JO TVNAUNOL 91S $9 “10a] Length (cm) Width (cm) Floral bract vestiture (abaxi- ally) Pedicel Color Vestiture Perianth Color (base/apex) Vestiture Length (cm Curvature, shape Staminode Color Apex Length (mm) up Length (cm) Width (cm) Flowering Habitat 8.5-11.5 6.5-9 Velutinous to vil- lous White to yellow Absent to puberu- lous White to yellow/ € Absent, to slightly puberulous api- 70-90°, parabolic White Cuspidate 6-8 1.7 1 Throughout year se secondary and rimary forest 12-15 7-9 Pubescent White to pink Absent to puberu- lous Pink/yellow Absent to puberu- lous 4.6-6 90°, parabolic White Cuspidate 7 (no fruits seen) (no fruits seen) January to August Open secondary growth 10-13 11-16 Absent to velutin- White to pink Absent White to pink/yel- ow Absent basally, ve- utinous apically 5-5.7 75-85°, sigmoid White, yellow at apex Acuminate 6-8 1.5-2 1-1.3 January to August Open secondary growth 6-11 8-12 Absent to villous White Absent to puberu- ous White/yellow suf- with pin Absent basally, pu- be scent apically and along sepal margins 70°, parabolic White, yellow at Acuminate 5-6 1.3-1.5 (January? to) Au- gust Open secondary growth 4.5-6.5 7.5-9 Absent to puberu- lous White to pink Absent to puberu- lous White/pink to red Absent basally, puberulous api- cally 3-4 50-70°, parabolic White Acute to acuminate 5-7 1-1.2 0.7-0.9 Throughout year Open secondary growth VINOOITAH ‘SSA [p86 LIS 518 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 65 Daniels and Stiles, 1979) was evaluated according to several criteria. In all cases the hybrid individual was intermediate in many of the parental mor- phological characters (see TABLE 8) and was easily distinguished in the field from both parents, which were always growing in the immediate vicinity. Except for Heliconia nutans x H. lankesteri, hybrids were usually found in areas of recent disturbance, indicating possible breakdown of habitat- and pollinator- isolating mechanisms. Hybrid pollen grains appeared distorted and highly vari- able in size under the scanning electron microscope (Kress & Stone, 1983), and they germinated poorly in sucrose solutions. Natural hybrids of Heliconia rarely produce viable seed (Stiles, 1979; pers. obs.). However, in the most common hybrid, H. mariae x H. pogonantha, mature fruits containing fully formed seeds have been found. In no case, except possibly in H. nutans x H. lankesteri, was any type of “hybrid swarm” apparent, all hybrids occurred as a single individual plant. Heliconia curtispatha Peterson x H. pogonantha var. holerythra Daniels & Stiles SPECIMENS EXAMINED. Panama. CoLon: along Rio Guanche, elev. ca. 100 ft, Kress et al. 80-1156 (DUKE). PANAMA: El Llano-Carti Rd., 15 km from Pan Am Hwy. turnoff, elev. 1200 ft, Kress & Hammel 80-1144 (DUKE). Heliconia mariae J. D. Hooker x H. pogonantha var. holerythra Daniels & Stiles SPECIMENS EXAMINED. Costa Rica. Limon: along road between Puerto Viejo and BriBri, elev. ca. 100 m, Kress et al. 76-627 (DUKE), Kress 77-795 (DUKE). Panama. CoLOn: along Rio Guanche, elev. 25 m, Kress 76-659 (DUKE); along roadside near Rio Buenaventura, 3 mi from Portobelo, ca. sea level, Kress 77-875 (DUKE). DARIéN: Marraganti and vic elev. 10-200 ft, Williams 686 (NY). Heliconia nutans Woodson x H. lankesteri var. rubra Daniels & Stiles SPECIMEN EXAMINED. Panama. CHIRIQUi: in forest along Rio Chiriqui Viejo below Cerro Punta, elev. 5500 ft, Kress & Clarkson 79-1089c (DUKE). Heliconia secunda R. R. Smith var. secunda x H. clinophila R. R. Smith SPECIMEN EXAMINED. Costa Rica. HEREDIA: near waterfall at Cinchona, elev. 4400 ft, Kress 77-777 (DUKE). Heliconia stilesii Kress x H. danielsiana Kress Reported by Daniels and Stiles (1979) to occur around Golfito, Puntarenas, Costa Rica. No specimens cited. Heliconia tortuosa Griggs x H. nutans Woodson Reported by Daniels and Stiles (1979) to occur at Las Alturas, Puntarenas, Costa Rica. No specimens cited. 1984] KRESS, HELICONIA 519 ACKNOWLEDGMENTS Many friends and colleagues have given freely of their time and advice during the preparation of this revision, and my thanks go to all. Dr. Timothy Plowman first suggested Heliconia as a possible genus in need of revision, and these pages are dedicated to him. Dr. Lindsay Clarkson’s unfading and unfailing moral and intellectual encouragement were indispen- sable to the completion of this work. Dr. Lennart Andersson has proved a venerable friend and critic during our lengthy debates about Heliconia. I look forward to our continued exchange of ideas. Drs. L. Andersson, J. Lundberg, D. Stone, R. White, R. Wilbur, and P. Stevens critically read and advised on various sections of the manuscript. I have had numerous discussions with Drs. F. G. Stiles, G. Daniels, L. Andersson, R. Wilbur, and D. Stone on the biology and taxonomy of Heliconia. Drs. C. Humphries, J. Lundberg, and L. McDade convinced me of the significance and importance of phylogenetic systematics. B. Hammel, L. McDade, J. Cooper- Smith, and S. Knapp helped with collections and in all kinds of weather endured my lengthy field notations. J. MacDougal, M. Turner, and B. Kirchoff helped solve problems seemingly small to them but important to me. W. L. Culberson shared his expertise in botanical Latin. S. Sellers, L. Eibest, and P. Atkins provided endless technical assistance in electron microscopy, manuscript prep- aration, and greenhouse care, respectively. S. Dickerson and J. Daniels typed the manuscript. During my stay in Costa Rica, Ms. Flor Torres and her staff at the Organization for Tropical Studies provided invaluable logistic support; at the Las Cruces Tropical Botanical Garden Robert and Katherine Wilson helped in numerous ways. I thank S. Schrohenloher, C. Taylor, and S. K. Hamilton for their artistic skill and botanical eye in the preparation of the illustrations. The expert and meticulous editorial assistance of E. Schmidt and S. Spong- berg was invaluable and enlightening. Financial assistance was provided by a Duke University Graduate School Research Award to W. J. Kress, by Biomedical Research Support Grant No. 303-3806 to D. E. Stone, and by Mr. Rob Montgomery, of Nahiku, Hawaii. The Stanley Smith Horticultural Trust provided funds in part for the publi- cation of the color plates. This material is also based upon research supported by the National Science Foundation (Grants DEB-7701556, DEB-7724612, and BSR-8306939) and is published with support of National Science Foundation Grant BSR-8317553. The Foundation provides awards for research and education in the sciences. The awardee is wholly responsible for the conduct of such research and prep- aration of the results for publication. The Foundation, therefore, does not assume responsibility for such findings or their interpretation. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author and do not necessarily reflect the view of the National Science Foundation. 520 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 LITERATURE CITED ABALO, J. E., & L. G. Moraes. 1982. Veinticinco (25) heliconias nuevas de Colombia. Phytologia 51: 1-61. _ & —._ 1983a. Doce (12) heliconias nuevas del Ecuador. Ibid. 52: 387- 413. & ———. 1983b. Diez (10) heliconias nuevas de Colombia. Ibid. 54: 411-433. Apanson, M. 1763. Gingembres. Zingiberes. Familles des plantes. Vol. 2. Pp. 61-67. Vincent, Paris. ANDERSSON, L. 1981. Revision of Heliconia sect. Heliconia (Musaceae). Nordic J. Bot. 1: 759-784. ArGENT, G. C. G. 1976. The wild bananas of Papua New Guinea. Notes Roy. Bot. Gard. Edinburgh 35: 77-114. ARISTEGUIETA, L. 1961. El género Heliconia in Venezuela. Instituto Botanico, Caracas. ARNOLD, E. N. 1981. oe phylogenies at low taxonomic levels. Z. Zool. Syst. Evolut. -forsch. 19: 1-35. BAKER, . G. 1893. eee of the genera and species of Museae. Ann. Bot. (London) 7: x rene H. S. 1971. Una nova especie de Heliconia L. (Musaceae) de le raque pendula. Rodriguésia 23: 127-130. 1976. Heliconiae novae Americanae (Heliconiaceae). [bid. 28: 129-132. _ 1978. Heliconia bahiensis Barreiros n. sp. (Heliconiaceae). Ibid. 29: 225-227. Bett, A. D., & P. B. ToMiinson. 1980. Adaptive architecture in rhizomatous plants. J. Linn. Soc., Bot. 80: 125-160. BENTHAM, G., & J. D. Hooker. 1880, Genera plantarum. Vol. 3. L. Reeve & Co.; Bisson, S., S. GUILLEMET, ine EIAMEL, 1968. Contribution al’étude caryo-taxinomique des Scitaminées. Mém. Mus. Natl. Hist. Nat., Sér. B, Bot. 18: 59-146. Bremer, K., & H.-E. WANNTORP. 1978. Phylogenetic systematics in botany. Taxon 27: 317-32 Coomss, E. nN K., M. J. Donocuue, & R. J. McGintey. 1981. Characters, computers, and cladograms: a review of the Berkeley cladistics workshop. Syst. Bot. 6: 359- 372. Cronguist, A. 1981. An integrated system of classification of flowering plants. Co- lumbia Univ. Press, New York. Daniaren, R. M. T., & H. T. Cuirrorp. 1982. The monocotyledons. A comparative study. Academic Press, New Yor & F.N. Rasmussen. 1983. Monocotyledon evolution. Sena and phylo- genetic estimation. Jn: M. K. HEcut, B. WALLAce, & G. T. PRANCE, eds., Evolu- tionary biology. Vol. 16. Pp. 255-395. Plenum Press, New Y on pa G.S. 1978. Lectotypification of Heliconia platystachys ances Kew Bull. 1, 682. EG. STILES. nae The Heliconia taxa of Costa Rica. Keys and descriptions. aS 15(Supl.): 1-15 Dopson, C. H., & A. H. ks 1978. Heliconias (Musaceae) of the Rio Palenque Science Center, Ecuador. Selbyana 2: 291-299. ELDREDGE, N., & J. CRACRAFT. 1980. Phylogenetic patterns and the evolutionary pro- cess. Columbia University Press, New Yor Emyaop10, L. pe MELLO Fitno. 1975. O género Heliconia na flora fluminensis de Frei ENGLER, A. 1900. Das Pflanzenreich. IV. 45. Musaceae (K. SCHUMANN). Wilhelm Engelmann, Leipzig 902. Das Src IV. 48. Marantaceae (K. SCHUMANN). Wilhelm En- gelmann, Leipzig. 904. Das Pflanzenreich. IV. 46. Zingiberaceae (K. SCHUMANN). Wilhelm En- gelmann, Leipzig. 1984] KRESS, HELICONIA 521 . 1912. Das Pflanzenreich. IV. 47. Cannaceae (Fr. KRANZLIN). Wilhelm Engel- mann, Leipzig. & K. PRANTL. 1889. Musaceae, Zingiberaceae, Cannaceae, pea Die natiirlichen Pflanzenfamilien, II. 6: 1-43. Wilhelm Engelmann, & 30. Musaceae, Cannaceae, Zingiberaceae, Mites Die na- tiirlichen Pflanzenfamilien, II. 15a: 505-693. Wilhelm Engelmann, Leipzig. ERDTMAN, G. 1966. Pollen morphology and plant taxonomy. pre & Wiksell, Stockholm. (Offset ena with addendum, Hafner, New Yor Farr, E. R., J. A. Leussi F. A. STAFLEU. 1979. Index nominnin genericorum. Farris, J. S. 1970. Methods for computing Wagner trees. Syst. Zool. 19: 83-92. GatTIN, M. C.-L. 1908. Recherches anatomiques sur |’ Be et la germination des Cannacées et des Musacées. Ann. Sci. Nat. Bot. 8: 113-146. GreEEN, P. 1969. Notes on Melanesian plants. II. Old ate Heliconia (Musaceae). Kew Bull. 23: 471-478. Gricocs, R. F. 1903. On some species of Heliconia. Bull. Torrey Bot. Club 30: 641- 664 Two es species of American wild bananas with a revision of the generic name. Ibid. 31: 4 hs 915. eae new species and varieties of Bihai. Ibid. 42: 315-330. HENnNIG, W. 1966. Phylogenetic systematics. (Translated by D. D. Davies & R. ZANGERL.) Univ. Illinois Press, Urbana. Humpnrey, J. E. 1896. The development of the seed in the Scitamineae. Ann. Bot. 10: 0. fe: HUTCHINSON, J. 1934. Families of flowering plants. ed. 1. Vol. 2. Monocotyledons. Clarendon Press, Oxford. 1959. Families of flowering plants. ed. 2. Vol. 2. Monocotyledons. Clarendon Press, Oxford. Kiuce, A.G., & J.S. Farris. 1969. Quantitative phyleti 1 the evolution of anuran Syst. Zool. 18: 1-32. Kress, W.J. 1981a. New Central American taxa of Heliconia (Heliconiaceae). J. Arnold Arbor. 62: 243-260. Systematics and reproductive biology of Central American Heliconia (Heliconiaceae). Unpubl. Ph.D. dissertation, Duke University, Durham, North Car- olina. . 1983a. Self-incompatibility in Central American Heliconia. Evolution 37: 735- 4 . 1983b. Crossability barriers in Neotropical Heliconia. Ann. Bot. (London) 52: 131-147. (in press). Pollination and reproductive biology of Heliconia. In: W.G. D’Arcy, ed., The natural history of Panama. Missouri Bot. Garden, St. Louis D.E. Stone. 1982. Nature of the a ea in mo onoco tyledons, with special reference to the pollen grains of Canna and Heliconia. Grana 21: 129-148. & 83. Morphology and eae significance of exine-less pollen of Heliconia (Heliconiaceae). Syst. Bot. 8: 14 7, ,&S. SELLERS. 1978. secede of exine-less pollen: Heliconia (Heliconiaceae). Amer. J. Bot. 65: 1064-1076. Kuntze, O. 1891. Revisio generum plantarum. Pars I. Arthur Felix, Leipzig Lang, I. E. 1955. Genera and generic relationships in Musaceae. Mitt. Bot. Siciseami. Miinchen 13: 114-131. LINHART, Y. B. 1973. Ecological and behavioral determinants = pollen dispersal in hummingbird-pollinated Heliconia. Amer. Naturalist 107: 511-523. Linnaeus, C. 1753. Species plantarum. Vol. 2. (1957 facsimile - ed. 1, Ray Society, 1771. Mantissa plantarum. (1961 facsimile reproduction, J. Cramer, Wein- a22 JOURNAL OF THE ARNOLD ARBORETUM [vVoL. 65 LunbsERG, J. G. 1972. Wagner networks and ancestors. Syst. Zool. 21: 398-413. 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Bot. 68: 49-6 TAKHTAJAN, A. L. 1980. Outline of the classification of flowering plants (Magnolio- phyta). Bot. Rev. (Lancaster) 46: 225-359. ToMLInSon, P. B. 1959. An anatomical approach to the classification of the Musaceae. Linn. Soc., Bot. 55: 779-809. . 1962. Phylogeny of the Scitamineae—morphological and anatomical consid- erations. Evolution 16: 192-213. Watrous, L. E., & Q. D. WHEELER. _ The out-group comparison method of character analysis. Syst. Zool. 30: Witey, E. O. 1978. The seers eee concept reconsidered. Syst. Zool. 28: 88— 1981. Phylogenetics—the theory and practice of phylogenetic systematics. J. ork. Woopson, R. E., Jr., & R WwW, ——- 1945. Flora of Panama. Musaceae. Ann. Missouri Bot. Gard. 32: 48-5 DEPARTMENT OF BOTANY Present address DUKE UNIVERSITY THE MARIE SELBY aa GARDENS DurRHAM, NorTH CAROLINA 27706 811 SoutH PALM SARASOTA, FLORIDA 33577 524 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 EXPLANATION OF PLATES PLATE I Three ee of the trichocarpa group. oe B, Heliconia necrobracteata: A, inflores- cence, x 0.2; B, flower, x |. C, D, H. m oe C, inflorescence, x 0.14; D, flower, x 0.8. E, F, H. talamancana: E. nha ca x 0.1; F, flower, x 0.8. PLATE II Two species of the trichocarpa group. A, C, Heliconia colgantea: A, inflorescence, x 0.1: C, flower, x 1. B, D, H. trichocarpa: B, inflorescence, x 0.1; D, flow l. PLATE III Three species of the nutans group. A, B, Heliconia marginata: A, inflorescence, x 0.25; B, flower, x 0.75. C, D, H. secunda: C, inflorescence, x 0.18; D, flower, x 0.75. E, F, H. nutans: E, inflorescence, x 0.25; F, flower, x 1. PLATE IV Two species of the nutans group. A, C, Heli l hys: A, inflorescence, x 0.08; C, flower, x 1.5. B, D, H. collinsiana var. collinsiana: B, inflorescence, x 0. 14; D, flower, x1. PLATE V Species of the curtispatha group. A, C, Heliconia curtispatha: A, inflorescence, 0.1; C, flower, x 0.9. B, D, H. stilesii: B, inflorescence, x 0.18; D, flower, x 0.8. PLATE VI Two species of the pogonantha group. A, C, Heliconia sessilis: A, inflorescence, x 0.13; C, flower, x 1.5. B, D, H. mariae: B, inflorescence, x 0.18; D, flower, x 1.5. PLATE VII Two species of the pogonantha group. A, C, Eas pogonantha var. veraguasensis: A, inflorescence, x 0.08; C, flower, x 0.75. , H. ramonensis var. lanuginosa: B, inflorescence, x 0.1; D, flower, x 1 PLATE VIII Three species of the pogonantha group. A, D, Heliconia danielsiana: A, inflores- cence, x 0.1; D, flower, x 0.8. B, E, H. magnifica: B, inflorescence, x 0.1; , flower, 8. C, F, a: xanthovillosa: C, inflorescence, x 0.1; F, flower, x 0.75. KRESS, HELICONIA PLATE I 526 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 PLATE II 1984] KRESS, HELICONIA a27 Ps PLATE III 528 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 , in : i 26st . a Fs i Ce PLATE IV 1984] KRESS, HELICONIA PLATE V JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 530 PLATE VI KRESS, HELICONIA 531 1984] PLATE VII JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 532 PLATE VIII 1984] ROSATTI, PLANTAGINACEAE 333 THE PLANTAGINACEAE IN THE SOUTHEASTERN UNITED STATES! THOMAS J. ROSATTI PLANTAGINACEAE A. L. de Jussieu, Gen. Pl. 89. 1789, ““Plantagines,”” nom. cons. (PLANTAIN FAMILY) A small family of terrestrial [to aquatic], mostly acaulescent herbs [or oc- casionally woody subshrubs] with simple, usually alternate, exstipulate leaves. Flowers mostly perfect, small, inconspicuous, usually numerous in cylindrical [to capitate] axillary spikes, mostly proterogynous and wind pollinated. Peri- anth for the most part actinomorphic and membranaceous-scarious. Sepals 4 [rarely 3], imbricate, the margins glabrous to [rarely long-]ciliate. Corolla sal- rform [or rarely the lobes indistinct] with 4 (sometimes 2) [rarely 1] stamens alternate with the petals, anthers versatile, filaments long and adnate to the tube. Gynoecium of 2 completely connate carpels, the style filiform, the ovary 'Prepared for the Generic Flora of the Southeastern United States, a long-term project currently of this area, with information about extraregional members of a family or genus in brackets [ ]. References I have not enhet are marked with an asterisk. I thank Norton Miller for the innumerable ways in which he contributed to the development of this treatment, and especially for the freedom he allowed in my pursuit of information regarding the Plantaginaceae. The expert help of Carroll Wood and George K. Rogers is very gratefully acknowl- ormation Speeecnan ang appreciated effort was put forth by Eleanor Gossen (New York State Museum), d University), and Jane Brennan and Lothian Lynas (New York Botanical ea Collections fom the fibaies and herbaria associated with these institutions were heavily relied upon throughout this study The Pane ta was prepared by Karen Stoutsenberger, under the direction of Carroll Wood and Kenneth R. Robertson, from fresh or preserved material rae by W ood and/or Robertson in Massachusetts and by Robertson and Ray E. Umber in North Carolin This mre is nablished as New York State Science Service J tal Series No. 433. © President and Fellows of Harvard College, 1984. Journal of the Arnold Arboretum 65: 533-562. October, 1984. 534 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 superior, 2- [rarely 1-]locular, with 1 to many anatropous to hemianatropous ovules per locule and axile [or basal or subbasal] placentation. Fruit a circum- scissile [or rarely indehiscent] capsule [or stony achene]. Seed with a longitu- dinally [rarely transversely] oriented embryo in fleshy endosperm, the seed coat more or less mucilaginous when wet. (Including Littorellaceae S. F. Gray; Psylliaceae Horaninow, nomen superfluum.) Type GeNus: Plantago L. Three genera and 220-275 species; represented in the southeastern United States by 12 (or perhaps 13) species of Plantago, a widely distributed and polytypic genus. Plantago differs from Bougueria Dene. and Littorella Berg. in its axile (vs. basal or subbasal) placentation, its two-locular (vs. one-locular) ovary, and its two- to many-seeded (rarely one-seeded), circumscissile capsule (vs. stony achene). Barnéoud divided the family into two tribes, primarily on the basis of these differences in fruit. The three species of Littorella (L. uniflora (L.) Ascherson, 2n = 24, of western and central Europe; L. americana Fern., 2n = 24, of northeastern North Amer- ica; and L. australis Griseb. ex Bentham & Hooker, of Chile and Argentina) occur on the shores of lakes and ponds and the margins of quiet streams. They are monoecious herbs with spicate inflorescences in which two carpellate flow- ers are nearly at ground level and are separated from a single, subterminal flower. (In L. australis the carpellate and staminate flowers may be more numerous and less separated.) The carpellate flowers develop one essentially sessile ovule, and at maturity the embryo is parallel to the long axis of the seed. In the single, slightly woody species of Bougueria (B. nubicola Dene., 2n = 12), a poorly collected plant of the high Andes in Peru, Bolivia, and Argentina, the inflorescence is a congested capitate spike in which perfect flowers are lower and more numerous than carpellate ones (although opinion varies on this point). The solitary ovules of both types of flowers are stalked, and at maturity the embryo is perpendicular to the long axis of the seed. The various proposed affinities of the Plantaginaceae have been much dis- puted, although several authors have suggested an alliance with the Plumba- ginaceae and/or Primulaceae, and in more recent years a relationship with the Scrophulariaceae has attained a certain level of acceptance. Endlicher united the Plantaginaceae and the Plumbaginaceae in his ““Plum- bagines”’ despite the fact that members of the Plumbaginaceae have five-merous flowers, stamens opposite the petals, and seeds with a mealy endosperm. De- caisne compared the Plantaginaceae with the Primulaceae, some members of which have circumscissile capsules, and Bentham and Hooker placed them near the end of their ““Gamopetalae,” calling it an anomalous family reminis- cent of the endospermous Acanthaceae in seed structure and insertion. The family was positioned between the Phrymaceae and the Rubiaceae in Die natitirlichen Pflanzenfamilien (see Harms & Reiche), and Bessey included it with the Primulaceae and the Plumbaginaceae in his Primulales. Hutchinson (1959) put the family in a monotypic order immediately after the Plumbagi- naceae (Primulales) and before the Crassulaceae (Saxifragales). Melchior po- sitioned an identically circumscribed Plantaginales between the Solanales and the Dipsacales (and the family between the Phrymaceae and Caprifoliaceae). 1984] ROSATTI, PLANTAGINACEAE 535 Airy Shaw (in Willis) proposed that the Plantaginaceae are allied with the Scrophulariaceae, stating that the “‘flower is usually regarded as derived from a five-merous type in the same way [1.e., by suppression of one calyx lobe and complete fusion of two corolla lobes] as that of Veronica (Scrophulariaceae), and there are good grounds (including entomological, viz. the food-plants of certain Lepidoptera and Coleoptera) for believing that the Plantaginaceae are in fact allied to Scrophulariaceae.’’ Most subsequent authors have conceded such a relationship. Thorne (1976) placed the family in his Bignoniales (with six other families) between the Scrophulariaceae and the Lentibulariaceae. Dahlgren included it in the Scrophulariales (composed of Thorne’s Bignoniales and ten other families). Takhtajan put it in the suborder Scrophulariineae between the Gesneriaceae and the Lentibulariaceae and stated that the Plan- taginaceae are ‘“‘near to and derived from Scrophulariaceae.” Cronquist (1981) more or less agreed with such an ancestry, but he nevertheless maintained the Plantaginales as a distinct order. (Although Cronquist earlier [1968] had noted that without phily and the persistent, scarious corolla, the Plantaginaceae could be positioned in the Polemoniales, he also pointed out that the occurrence in the family of both micropylar and chalazal haustoria was in agreement with a relationship to the Scrophulariaceae.) Numerous studies have indicated that the Plantaginaceae are rather ho- mogeneous chemically. Hegnauer discovered that species of both Plantago and Littorella produce aucubin (an iridoid glucoside; Jensen et a/. indicated a com- mon origin for all iridoid-containing taxa), and Bourdu et a/. showed that species of these two genera store the same saccharides (stachyose being the most abundant) in comparable proportions in the roots and stems. The seeds of L. uniflora contain planteose and saccharose; those of Plantago have plan- teose but in some species stachyose as well (Bourdu et a/., Bourdu & Gorenflot). Wallaart identified sorbitol (a sugar alcohol) from the leaves of L. uniflora and ten species of Plantago and considered the accumulation of this hexitol to be a family character. Aucubin and planteose were also identified in the seeds of eight species of Plantago (including P. major L.) in Egypt (Ahmed et a/.). Chemical evidence has also suggested that the Plantaginaceae are much more closely related to the Scrophulariaceae than to either the Plumbaginaceae or the Primulaceae (Bourdu et a/., Gibbs, Hegnauer). Aucubin has been found as well in the Globulariaceae, Loganiaceae, Orobanchaceae, and Scrophulariaceae (Bate-Smith). The production of mannitol (a stereoisomer of sorbitol) by two species of Plantago also indicates a strong biochemical link with the Scroph- ulariaceae, according to Wallaart. The Plumbaginaceae and Primulaceae are among 68 families in which both leuco-anthocyanins and trihydroxy constit- uents have been identified, while the Scrophulariaceae and Plantaginaceae are among 70 families in which these compounds are not known (Bate-Smith). Seigler reported that while members of the Plumbaginaceae and Primulaceae do not have iridoid monoterpenes, such compounds have been found in about 41 other families, including the Scrophulariaceae and the Plantaginaceae. Ac- cording to Alston and Turner, both the Scrophulariaceae and the eer al are among about 20 families, exclusive of the Plumbaginaceae and the Prim- ulaceae, in which linoleic acid and oleic acid are the major fatty acids. 536 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 The most common and widespread (and therefore, presumably, the ancestral) base chromosome number in the Plantaginaceae is six. (The Scrophulariaceae have six among their base numbers, but only the Plumbaginaceae have chro- mosomes with any morphological similarity to Plantago [McCullagh].) Dip- loids, triploids, tetraploids, hexaploids, octaploids, 12-ploids, and 16-ploids (i.e., 2n = 12, 18, 24, 36, 48, 72, and 96) are known in Plantago, while only diploids and tetraploids (in the same series) have been reported for Bougueria (Rahn) and Littorella, respectively. The base numbers five (diploids, tetra- ploids, hexaploids, and decaploids) and four (diploids and tetraploids) have numerous levels have also been reported in Plantago (see primarily Bolkhov- skikh et al.). Species of Plantago and Littorella shed binucleate pollen, except that that of P. lanceolata L. is trinucleate (Brewbaker). Bougueria 1s evidently unknown in this regard. The pollen in the Plantaginaceae (which provides no clear indication of the systematic placement of the family [Erdtman]) is usually spheroidal (but some- times ovoid or irregular) and more or less distinctly periporate, with the pores circular or irregular in shape. After some initial confusion that appears to have been resolved by the use of scanning electron microscopy, there is now agree- ment that the sculpturing is more or less verrucate. Although the outermost layer of ornamentation has been alternately described as micro-echinate (Bas- sett ef al., Bassett & Crompton) or scabrate (Clarke & Jones), the latter is the more appropriate term (according to Clarke & Jones) because the projections are rounded. Discrepancies regarding the presence or absence of annuli and opercula also exist in the literature, apparently because of a failure to recognize that within the family a continuum exists between grains possessing and those completely lacking such features (Clarke & Jones). Because only four species of the Plantaginaceae are in genera other than Plantago, studies of topics such as embryology, anatomy, and morphology have mostly involved only species of Plantago and are therefore summarized in the generic discussion. REFERENCES: AHMED, Z. F., A. M. Rizk, & F. M. HAMMouDA. Phytochemical studies of Egyptian Plantago species (glucides). Jour. Pharm. Sci. 54: 1060-1062. 1965. ALSTON, R. E., & B. L. TURNER. Biochemical systematics. xii + 404 pp. Englewood ARBER, A. Water plants[.] A study of aquatic angiosperms. Frontisp. + xvi + 436 pp. Cambridge. 1920. [Various references to Plantago and Littorella Baivey, L. H., E. Z. BAttey, & BAiLtEy HortTorium STAFF. Hortus third. xiv + 1290 p. New York and London. 1976. [Littorella, 673, sometimes grown in aquaria; Plantago, 882, several species cultivated for ‘‘psyllium” seed, which is used as a mild laxative.] BaiLLon, H. Plantaginacées. Hist. Pl. 9: 274-280. 1888. BaRNEOuD, F. M. Mémoire de botanique. Recherches sur le développement, la structure générale et la classification des Plantaginées et des Plumbaginées. Mémoire de Géo- 1984] ROSATTI, PLANTAGINACEAE 537 logie. De l’origine des lacs. Théses pour le doctorat. Acad. Paris Fac. Sci. 44 pp. 2 pls. Paris. 1844. [Plantaginées, 1-19, 42, 43, pl. J; 10 sections and 114 species of Plantago recognized, species merely listed.] Monographie générale de la famille des Plantaginées. 52 pp. Paris. 1845. [Same sections and species of Plantago recognized in 1844, but species more fully treated.] BAssETT, I. J., & C. W. Crompton. Pollen morphology and chromosome numbers of the family Plantaginaceae in North America. Canad. Jour. Bot. 46: 349-361. 1968. [29 spp. (including five subspp. and one var.) of Plantago, including 12 (or perhaps 13) present in the Southeast, and Littorella americana; light micrographs of most, key included.] , & J. A. PARMELEE. An atlas of airborne pollen grains ae common fungus spores of ‘Canada. 321 pp. Research Branch, Canada Dep. Agr. Ottawa. 1978. [Plantaginaceae, 216-226; seven spp. of Plantago and Littorella americana; light and scanning elecron micrographs. ] BATE-Smitu, E. C. The phenolic constituents of plants and their taxonomic significance. I. Dicotyledons. Jour. Linn. Soc. Bot. 58: 95-173. 1962. [Plantaginaceae, aucubin, 114, 159. BENTHAM, G., & J. D. Hooker. Plantagineae. Gen. Pl. 2(2): 1223-1225. 1876. Bessey, C. E. Phylogenetic taxonomy of flowering plants. Ann. Missouri Bot. Gard. 2: 109-164. 1915. [Plantaginaceae, 143.] BOLKHOVSKIKH, Z., V. GriF, T. MATVEJEVA, & O. ZAKHARYEVA. Chromosome numbers of flowering plants. A. A. Feporov, ed. oo and English prefaces.) 926 pp. Leningrad. 1969. eee pane 488-4 Bourpbu, R., D. CARTIER, & R. GORENFLOT. (cae biochimiques des genres Littorella et Plantago. Bull. Soc. Bot. prance 110: 107-109. p/. J. 1963. R. GorenFLot. Sur les galactosides des graines dans le genre Plantago L. Compt. Rend. Acad. Sci. Paris 253: 698-700. 1961. [Seeds of species in 11 sections (sensu Pilger); all contained planteose; stachyose occurred only in species of sects. Arnoglossum Dene. and Hymenopsyllium Pilger.] BREWBAKER, J. L. The asibution and phylogenetic oe of binucleate and trinucleate pollen grains in the angiosperms. Am. Jour. Bot. 54: 1069-1083. 1967. Casper, S. J. Plantaginaceae. /n: G. HEI, Illus. F1. ieee ed. 2. 6(1): 559-608. 1 974, CiarkE, G. C. S., & M. R. Jones. The northwest European pollen flora, 15[.] Planta- ginaceae. Rev. Palaeobot. Palynol. 24: 129-154. 1977. Corner, E. J. H. The seeds of dicotyledons. Vol. 1. xii + 311 pp. Cambridge, London, New York, and Melbourne. 1976. [Plantaginaceae, 218; an “alliance with the uni- tegmic Polemoniaceae and Scrophulariaceae accords better with the simple seed- structure than with the bitegmic Primulaceae.”’] Cronauist, A. The evolution and classification of flowering plants. xi + 396 pp. Boston. 1968. [A unifamilial Plantaginales positioned between the Lamiales and the Scroph- ulariales in the Asteridae.] An integrated system of classification of flowering plants. Frontisp. + xvili + 1262 pp. New York. 1981. [The Plantaginaceae put in its own order between the Hydrostachyaceae (Callitrichales) and the Scrophulariaceae (Scrophulariales) in the Asteridae; pollen assignable Plantago from the middle or late Miocene; see also Nordic Jour. Bot. 3: 75-83. DAHLGREN, R. M. T. A revised mite of classification of the angiosperms. Bot. Jour. Linn. Soc. 80: 91-124. 1980. , S. ROSENDAL-JENSEN, & B. J. NIELSEN. A revised system of classification of the a giosperms with comments on correlation between chemical and other characters. Pp. 149-204 in D. A. YounG & D. S. SEIGLER, eds., Phytochemistry and angiosperm phylogeny. New York. 1981. [Placement of the Hippuridaceae near the Plantagi- naceae may be indicated on chemical grounds; see also Jensen et al.] 538 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Davis, G. L. Systematic embryology of the angiosperms. x + 528 pp. New York, Lon- don, and Sydney. 1966. bea eae 212, but only P/antago actually mentioned. ] DECAISNE, J. Plantaginaceae. DC. Prodr. 13: 693-737. 1852. [17 sections of Plantago.] DietricH, H. Cytologische Cae na innerhalb der Familie der Plantaginaceae I[.] Chromosomenzahlen einheimischer Arten der Gattungen Plantago und Litto- rella. Wiss. Zeitschr. Friedrich Schiller Univ. Jena Math. Nat. 21: 945-951. 1972.* II[.] Zusammenstellung bekannter Chromosomenzahlen und der Vergleich zwischen Grundzahl und Lebensdauer. Jbid. 24: 437-461. 1975.* III[.] Cytotaxonomische Ergebnisse. (Summaries in German, English, Russian, and Spanish.) [bid. 29: 559- 587. 1980. [Includes chromosome counts reported 1975-1979; author’s counts for 36 species, subspp., and vars. of Plantago, six species of Psyllium, and Littorella uniflora, Plantago fon Micropsyllium Dene. emend. H. Dietr. and Diandrae H. Dietr., sect. nov.; six new combinations under Psy//ium Miller. Presumed base chro- mosome number i is six, from which are derived five and four.] EICHLER, A. W. Bltithendiagramme. Erster Theil. 348 pp. Leipzig. 1875. [Plantaginaceae, 24-288.] ENDLICHER, S. L. Plantagineae. Gen. Pl. 5: 346-348. 1837. ERDTMAN, G. Pollen morphology and plant taxonomy. Angiosperms. Frontisp. + xii + 539 pp. Uppsala. 1952. [Pollen of Bougueria nubicola described, 324.] FERNANDES, A., & F. FRANCA. Contribution a la connaissance cytotaxinomique des Spermatophyta du Portugal VI. Plantaginaceae. Bol. Soc. Broter. I. 46: 465-501. London. 1974. [Vol. 4 includes bibliography, index, and addendum; numerous references to the Plantaginaceae. ] GLEASON, H. A., & A. CRonquist. Manual of vascular plants of northeastern United States and adjacent Canada. 1i + 810 pp. New York, Cincinnati, Toronto, London, and Melbourne. 1963. [Plantaginaceae, 642-644.] Gopwin, H. The history of the British flora. ed. 2. x + 541 pp. 28 pls. Cambridge, London, New York, and Melbourne. 1975. [Plantaginaceae, 326-332; pollen record in Britain indicates a progressive decline in Littorella uniflora since last glaciation (seen as result of natural infilling of sane lakes), as well as an ancient natural status for Plantago lanceolata and P. major.] Harms, H., & C. ReicHe. Plantaginaceae. Nat Pflanzenfam. IV. 3b: 363-373. 1895. [Treatment of Plantago by Harms only; subg. Plantago, 11 sections; subg. Psyllium established. ] HEGNAUER, R. Chemical evidence for the classification of some plant taxa. Pp. 121- pentacyclic triterpene carboxylic acids, stachyose-type oligosaccharides, hexitols, planteose, fatty oil, and reserve celluloses in the Plantaginaceae.] . Koorman. Die systematische Bedeutung von iridoiden Inhaltsstoffen im Rahmen von Wettstein’ s Tubiflorae. Pi. Med. 33: l= 33.1978.[A catapol well as aucubin and catapol, awe in Plantaginaceae; a relationship to Tubiflorae, perhaps Scrophulariaceae tribe Veroniceae indicated. Melittoside noted in | yoideae; aucubin and catapol absent in Labiatae. Melittoside not reported from pee see Swiatek ef a HUTCHINSON, J. e families of flowering ae ed. 2. Vol. 1. Dicotyledons. xv + 510 pp. Oxford. ee Siarananrr 455, 456. . Evolution and phylogeny of flowering plants. xxv + 717 pp. London and New York. 1969. aden $59-562.] . The families of flowering plants. ed. 3. xx + 968 pp. Oxford. 1973. [Plantagi- naceae, 30, 149, 561-564.] 1984] ROSATTI, PLANTAGINACEAE D359 JENSEN, S. R., B. J. NIELSEN, & R. DAHLGREN. Iridoid compounds, their occurrence and systematic importance in the angiosperms. Bot. Not. 128: 148-180. 1975. [Aucubin and its derivative catalpol, both Group I iridoids (10-hydroxylated), indicated for Plantago (cf., however, Swiatek et al. regarding melittoside, another iridoid possibly e€ puridales (with Group I iridoids) perhaps near the Scrophulariaceae or Plantagi- naceae. ] McCutacu, D. Chromosomes and chromosome morphology in Plantaginaceae I. Ge- netica 16: 1-44. 1934 Metcuior, H. Reihe Plantaginales. Jn: H. MELCHIOR, Engler’s Syllabus der Pflanzen- familien, ed. 12. Gs 471, 472. 1964. Moncontié, C. Les stomates des earns tas (English summary.) Revue Gén. Bot. 76: 491-529. ee. [Littorella uniflora; 18 spp. of Plantago Moore, D. M., ed. Plantaginaceae. Jn: T. G. TUTIN ef al., eds., FL. Europaea 4: 38-44. 1976. [Plantago by A. O. me oe & D. CARTIER; Littorella by D. M. Moore.] Plantaginaceae. P. 2411 V.H. HeEyYwoop, ed., Flowering plants of the world flowers on the same plant—cf., however, Cronquist (1981) and Harms & Reiche, who state that the genus is Se tate (although opinion seems varied on this point).] MULLER, J. Fossil pollen records of extant angiosperms. Bot. Rev. 47: 1-142. 1981. [Plantaginaceae, 97, 98; pollen generally listed in pre- -Quaternary records as either Plantaginacearumpollis or Plantago may include that of Littorella.] aes R. Plantaginaceae. Pflanzenr. IV. 269(Heft 102): 1-466. 1937. [Two subgenera 19 sections of Plantago.] ret K. In: A. Léve, ed., IOPB chromosome number reports LHI. Taxon 25: 483- 500. 1976. Ui ac aaaaa 499, 500; Bougueria nubicola, 2n = 12; counts in 28 species of Plantag Scoain, R. Amino acid sequence studies and plant phylogeny. Pp. 19-42 in D. A. Younc & D. S. SEIGLER, eds., Phytochemistry and angiosperm phylogeny. New York. 1981. [In an “affinity tree” of angiosperms based on partial (in most cases 40 of 99 amino acid residues) sequence data regarding ieee cs from 38 en Plantago major is closest to Verbascum Thapsus, Digitalis purpurea, and Antirrhi- num majus (Scrophulariaceae), with only a few exceptions, genera within ‘ ves families are clustered toget SEIGLER, D. S. Terpenes and plant phylogeny. Pp. 117-148 in D. A. Younc & D. S. SEIGLER, eds., Phytochemistry and angiosperm phylogeny. New York. 1981. Swarprick, J. T., & J. C. RayMonp. The identification of the seeds and achenes of the British Plantaginaceae. Ann. Bot. II. 34: 1123-1129. pls. 1, 2. 1970. [Seeds of P. lanceolata and P. major, achenes of Littorella aie SwIATEK, L., D. LEHMANN, R. K. CHAUDHURI, & TICHER. Occurrence of melittoside in the seeds of Plantago media. ie te io 20: 2023, 2024. 1981. [Refer to Hegnauer and Kooiman as recording melittoside in Labiatae and Scrophulariaceae, but those authors record melittoside only from Labiatae, not Scrophulariaceae. Also TAKHTAJAN, A. L - Outlin of the classification of flowering plants (Magnoliophyta). Bot. Rev. 46: 225- 359. 1980. ee 297.] TuHorne, R. F. A phylogenetic anne of the Angiospermae. Evol. Biol. 9: 35- 106. 1976. [Plantaginaceae, 63; see also Thorne, 1983. Proposed new ea in re angiosperms. Nordic oe Bot. 3: 85-117. 1983. [Position of Plantaginaceae unchanged relative to Thorn WALLAART, R. A. M. Acyclic polyols as taxonomic characters. IL] Sorbitol in Plan- taginaceae and joint occurrence of sorbitol and mannitol in some Plantago species. Proc. Nederl. Akad. Wet. C. 84(1): 77-82. 1981. [The joint occurrence of two 540 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 stereoisomeric hexitols in one higher plant species is reported for the first time (in two spp. of Plantago).] Wiis, J.C. A dictionary of the flowering plants and ferns. ed. 7. Revised by H. Airy SHAW. xxii + 1214 + lili pp. London. 1966. [Plantaginaceae, Plantago, an itorella 661, 662.] Youna, D. J., & L. Watson. The classification of dicotyledons: a study of the upper levels of the hierarchy. Austral. Jour. Bot. 18: 387-433. 1970. [Data on 83 attributes of 543 genera; Plantago has characteristics of both “‘asclepioids” and ‘“‘acanthoids” (but more in common overall with the latter, which includes the Scrophulariaceae and six other families). ] 1. Plantago Linnaeus, Sp. Pl. 1: 112. 1753; Gen. Pl. ed. 5. 52. 1754. Erect or suberect, acaulescent (but sometimes branched near the base and/ or rhizomatous) [or caulescent (and branched)], terrestrial, annual, biennial, or perennial herbs [or occasionally woody subshrubs]. Leaves simple, linear, lan- ceolate to oblanceolate, spatulate, ovate, or cordate; alternate, in basal rosettes [or, less frequently, alternate or opposite on sparsely to densely branched stems], exstipulate, with dilated, sometimes sheathing bases, glabrous or (rarely) dense- ly pubescent (at least adaxially), entire, dentate, [or pinnatifid], one to many nerved, the blade and petiole distinct or not. Plants with only perfect flowers (or otherwise, usually with perfect and carpellate flowers on separate plants) [rarely dioecious]. Inflorescences axillary with cleistogamous or chasmogamous flowers sparsely to densely organized into long to short, cylindrical [to capitate] spikes. Flowers single in the axils of bracts, small, usually numerous, (chas- mogamous flowers) mostly proterogynous and wind pollinated, and actino- morphic or slightly zygomorphic (with respect to corolla lobes). Sepals 4, im- bricate, membranaceous-scarious, equal or in 2 unequal pairs (the abaxial pair sometimes connate in the basal half), alternate with the subtending bract, and often keeled [or rarely with many distinct nerves]. Corolla salverform with the tube ovoid to cylindrical and the 4 erect, spreading, or recurved lobes distinct, imbricate, membranaceous-scarious, and alternate with the sepals. Stamens 4 (occasionally 2), adnate to the corolla tube (rarely below the gynoecium), al- ternate with the petals; filaments filiform, induplicate in bud; anthers included or exserted, cordate, often apiculate, 2-locular, dorsifixed, versatile, and with longitudinal, latrorse to slightly introrse dehiscence. Carpels 2, connate; stigmas included or exserted; style filiform; ovary superior, 2-locular [or imperfectly Ficure 1. Plantago.a-j, P. /anceolata: a, habit, inflorescences with flowers immature, or with only styles protruding, or at anthesis, x '2; b, flower bud subtended by bract, x 6; c, flower with receptive style protruding, subtending bract in front, x 6; d, flower, later stage with corolla lobes open and anthers dehiscing, x 6; e, versatile anther, x 12; f, portion of corolla tube 1 section, filament bases inserted between folds of corolla, x 12; g, gynoecium, x 1 , capsule see by persistent calyx, x 6; i, adaxial surface of moistened seed, outer layer mucilaginous, = 6; j, embryo, x 6. k-m, P. aris- tata: k, habit—note long inflorescence bracts, x '4; 1, flower at anthesis, x 6; m, “‘cleis- 1984] ROSATTI, PLANTAGINACEAE 541 togamous” flower, x 6. n, 0, P. Rugelii: n, habit, x 2; 0, fruit enclosed in persistent corolla tube, x 6. p, q, P. major. p, fruit with circumscissile line of dehiscence, x 6; q, adaxial surface of seed, x 12. 542 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 3- or 4-locular by the intrusion of false septa from the carpel midrib]. Ovules 1 to many per locule, tenuinucellar, hemianatropous to anatropous, integument single, massive; placentation axile. ae a globose to elongate capsule, circum- scissile at or below the middle [ tin P. macrocarpa Cham. & Schlecht.]; calyx persistent, the corolla usually so (but then irregularly torn about the middle of the tube). Seeds yellow-tan to almost black, often shining, for the most part narrowly elliptic in longitudinal median section (through the widest dimension of the seed); seed coat more or less mucilaginous when wet; en- dosperm cellular, fleshy; embryo parallel to the long axis of the seed and straight or sometimes curved. (Including Coronopus Miller, non Zinn, nom. cons., and Psyllium Miller.) Lecrotyre species: P. major L.; see Britton & Brown, Illus. Fl. No. U.S. & Canada, ed. 2. 3: 245. 1913. (Plantago listed by Linnaeus [Critica Botanica, p. 110. 1737; p. 87 in A. Hort, English transl., 1938] as a Latin name given by the “Ancients” [Pliny and other Roman writers]. According to Clute, and most others who have commented on the issue, the name is from Latin planta, signifying the sole of the foot, and ago, used in plant names in the sense of “wort” [but also to mean having the characteristics of].)— PLANTAIN. A nearly cosmopolitan genus occurring largely in the Temperate Zones but only poorly represented in the lowland tropics, Plantago is recognized in the most recent worldwide monograph (Pilger) to consist of 258 species. Although perhaps best known for its nearly ubiquitous weedy members (P. /anceolata L. and P. major L., for example, are extremely widespread and even occur at low elevations in the New World tropics [D’Arcy; Dietrich, 1979]), most species are in fact of much more restricted distribution. Some are endemic to very small areas and/or are highly specialized ecologically. The genus is best rep- resented in the Mediterranean region, the Himalayas, southwestern North America, and the mountains of South America. Since Linnaeus (Species Plantarum, 1753) grouped the species of Plantago into those with scapo nudo and those with caule ramosa, suprasectional taxa have been recognized in the genus by Barnéoud (1844; two units based on seed number per capsule and seed shape in transverse median section), Harms (in Harms & Reiche; two subgenera based on leaf arrangement), Pilger (two sub- genera based on leaf arrangement), and Rahn (1978a; three subgenera based on oOvule/seed number per ovary/capsule, seed shave in transverse median section, corolla-tube pubescence, leaf shape, embryo orientation [plane between the cotyledons either parallel, oblique, or perpendicular to the adaxial surface of the seed], and chromosome number). The genus has also been divided into various numbers of sections by Barnéoud (1844), Decaisne, Harms (in Harms & Reiche), Pilger, and Rahn (1978a) based on different combinations of char- acters including spike morphology (size, shape, and flower arrangement and density), corolla pubescence, corolla-lobe shape, anther and stigma inclusion or exsertion, seed size, seed number per capsule, seed shape (in transverse median section as well as longitudinal median section through the widest di- mension of the seed), leaf shape, degree of stem lignification, plant duration, and geographic distribution. Rahn (1978a) recently typified a number of subge- 1984] ROSATTI, PLANTAGINACEAE 543 neric and sectional names in Plantago, and his infrageneric classification is followed here. Approximately 24 indigenous and eight naturalized species of Plantago rep- resenting all three subgenera and ten of the sections enumerated by Rahn (1978a) occur in North America. Twelve (or perhaps 13) species in two sub- genera and six (or perhaps seven) sections grow in the southeastern United States. The first four sections discussed below were placed in subg. PLANTAGO; the last three in subg. PsyLL1um (Miller) Harms in Harms and Reiche. Ten of our species are indigenous to North America, and four of these appear to be naturalized in the Southeast (i.e., are native to other parts of the continent). The predominantly Old World sect. PLANTAGO (fibrous-rooted perennials with petiolate leaves, elongate-cylindrical spikes, exserted anthers and stigmas, and 4—20 seeds per capsule) comprises about 19 species, including a closely related pair occurring in the Southeast. Plantago major, 2n = 12, 18, 24, thought to be a native of Europe and northern and central Asia but now established throughout most of the rest of the world, is a weed that reportedly occurs in most parts of our range, although it is poorly collected and/or perhaps rare in our area. Plantago Rugelii Dcne., 2n = 24, originally native to eastern North America, is a weedy species now widely established westward but at best ncommon outside of this continent. (It grows at or near sea level in Cuba [Dietrich, 1979] and has evidently become established in northern Europe [Hansen].) Plantago Rugelii occurs throughout the Southeast, but it seems to be poorly represented, if not absent, in peninsular Florida. Plantago Rugelii differs from P. major in its more elongate capsules that dehisce closer to the base and often contain fewer seeds; otherwise the species are remarkably similar morphologically. Although P. major and P. Rugelii usually grow (but not nec- essarily together) in fields, lawns, roadsides, sidewalks, and paths, both Tessene (1968) and Bassett (1973) have reported that P. major does not thrive in shaded and/or continually wet places and that P. Ruge/ii may be found in open woods and along damp shorelines. Hamilton and Buchholtz, however, counted sig- nificantly fewer seedlings of P. Rugelii in shaded plots than in open ones (all plots naturally seeded). Plantago major may be particularly well adapted to areas where the soil has been trampled (Blom; Sagar & Harper, 1960; Warwick) or compacted (Blom; see, however, Noe & Blom and Tessene, 1968). Hawthorn and Cavers (1976) found that in natural (pasture) populations, P. Rugelii had a longer half-life than P. major with regard to both seedlings and mature plants, while individuals of P. major flowered in greater numbers (proportionately) and earlier in development. Several authors have reported that P. major can be either an annual or a perennial (see Hawthorn & Cavers, 1976). Four of the six species comprising the Old and New World sect. MICROPSYLLIUM Dene. (taprooted annuals with linear leaves and loose, elongate spikes bearing small flowers with usually two stamens and included anthers and stigmas) occur in North America (see Bassett, 1966), and the two present in our range, Plantago heterophylla Nutt., 2n = 12, 22, and P. pusilla Nutt., 2n = 12, are nearly indistinguishable. Both have erect corolla lobes, but P. heterophylla has many (usually 10-25) small seeds per capsule, while the fruits 544 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 of P. pusilla almost invariably contain four larger seeds.? Very small individuals occur in both species. Plantago heterophylla is more common than P. pusilla in the Southeast (in North America it is almost restricted to our area). It grows in moist, sandy or silty soils and is frequently a weed in cultivated or waste places. It is an adventive in California (Pilger) and in the northeastern United States (northward to southern New York [Bassett, 1966]), but it is evidently indigenous in Argentina and Paraguay (Pilger). Plantago pusilla is most often found in disturbed areas. Its rather limited natural range includes the southern ton (Bassett, 1966). In our area, therefore, it occurs as a native in Arkansas and Tennessee and as an adventive in northeastern North Carolina The nearly cosmopolitan and rather poorly distinguished sect. PALAEOPSYLLIUM Pilger (thick-rooted perennials with ovate-cordate to oblan- ceolate leaves having relatively distinct petioles; loose, elongate spikes bearing large flowers with exserted anthers and stigmas; and two to nine seeds per capsule) consists of about 28 species, of which four are North American (see Bassett, 1967). Two of these occur in the Southeast. Plantago cordata Lam., 2n = 24, has distinctly petiolate leaves with ovate to cordate blades and veins that appear to arise from a midrib (vs. from the base of the blade, as in the rest of the genus), hollow scapes, sepals that are about as long as the bracts, and two- to four-seeded capsules that dehisce approximately at the middle. The species is unique in the genus in that the fruits are still green and alive at the time of dehiscence, when the lid of the pyxis readily falls off, and the two or three seeds with the entire fleshy placenta fall out as a unit (Tessene, 1969). This structure is buoyant and may represent an adaptation to dispersal by water. The plants grow in alluvial woods, on wet rocky banks, in streams through areas of slate bedrock, and in the shallow soil of cedar (Juniperus virginiana) barrens in our area. The species probably occurs in all of the southeastern states, although it is perhaps absent from southern Florida and is apparently unknown outside of North America. Plantago oe differs from P. major and P. Rugelii, with which it is often confused, in number of ways. Its leaves are larger, the blades are more abruptly patie to the petiole (but not always cordate), and the major veins, although actually parallel, approach a pinnate arrangement. In addition, the roots are thicker and the flowering time is earlier in P. cordata. 2Plantago laeti Saas Nutt. (1835; see Foster) has been the most commonly used name for the plants with ten or more seeds per capsule. Godfrey and Wooten used the earlier name P. hybrida W. Barton (1818) ee this species and incorrectly listed P. elongata Nutt. (instead of P. at ate in synonymy. The species with four seeds per capsule has been called P. elongata Pursh (181 pusilla Nutt. (1818), depending on taxonomic interpretation, the later name being used if our eastern plants are considered to be specifically distinct from more western populations. Tare os seed number is mentioned in none of the four eae es involved, and I have not been able to study the type specimens. To minimize the likelihood of adding confusion, I have chosen to use the most commonly employed names, P. eee and P. pusilla, although the reader should be aware that the situation is subject to chan 1984] ROSATTI, PLANTAGINACEAE 545 The second species, Plantago sparsiflora Michaux, 2n = 24, has oblanceolate leaf blades that gradually taper to the petiole, scapes that are only sometimes hollow, sepals that are about twice as long as the bracts, and two-seeded capsules that dehisce below the middle. It is known only from the Coastal Plain of northern Florida and the southeastern portions of Georgia and the Carolinas, where it is frequent in wet or dry savannas, roadsides, and ditches. Tessene (1969) studied in detail many aspects of Plantago cordata, which he considered to be the rarest North American species of the genus. He deter- mined that the characteristic venation of the leaf, so frequently used to distin- guish the species, usually does not develop until after fruiting has begun, and that depauperate inflorescences often retain solid peduncles. He stated that it usually grows in calcareous substrates and reported only one instance in which the species was found in a disturbed, man-made habitat (an unshaded drainage ditch). Tessene also attempted crosses between P. cordata and eight other species in four sections of Plantago, with only one F, plant resulting. This hybrid, P. cordata x P. eriopoda Torrey (both of sect. PALAEOPSYLLIUM) was sterile and intermediate in most characters. The New World sect. Vircinica Barn. (leaves elliptic to spatulate, often dentate; scapes long and with a terminal, many-flowered, cylindrical spike; abaxial sepals much narrower than the adaxial ones; corolla tubes glabrous and lobes erect or spreading; capsules with up to four but usually two or three seeds) was the subject of an ambitious revision based primarily on the morphology of herbarium specimens but influenced by ecological data, cultivation exper- iments, chromosome studies, hybridizations, and taximetric analyses (Rahn, 1974). Included were sections OREOPHYTUM Dene., CLEIOSANTHA Dene., and Novorsis Dene., in addition to species from sections PALAEOPSYLLIUM and LEUCOPSYLLIUM Dene. Twenty-eight species (four described as new) were rec- ognized in the section (morphologically indistinguishable entities differing in chromosome number were treated as conspecific); five of these occur in North America, and two are found in the Southeast. Both have concave adaxial seed surfaces (vs. plane in the three other species) and are annuals with slender primary roots, pilose leaves, spreading hairs on the scape, erect corolla lobes, and two-seeded capsules. Plantago virginica L., 2n = 12, 24 (orca. 24 [Rahn, 1974]), occurs throughout the eastern half of the United States, including all of the Southeast, and is apparently a waif in southern Ontario (Bassett, 1973). It is more sporadically distributed in the western states and northeastern Mexico. The species appears to be a spring ephemeral (Rahn, 1974; all but four of the 111 specimens I examined were collected earlier than June) and is generally found in dry, sandy soils of waste places, fallow and cultivated fields, and roadsides. Plantago virginica can be distinguished from the other common plantagos in our area by its dentate, spatulate leaves, its rather dense, cylindrical spikes, and its persistent corollas with conspicuous, yellow, erect lobes. Depauperate forms of the species are fairly common. Plantago rhodosperma Dene., 2n = 24, occurs in rocky or sandy soils mainly in the southwestern United States and adjacent Mexico. Although Rahn (1974) cited no specimens from our range (and I have seen none), his distribution 546 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 map for the species includes Louisiana, most of Arkansas and Mississippi, and southwestern Tennessee. Several other authors (Correll; Small, 1913; Smith; Steyermark) have reported the species from various combinations of these states. Plantago rhodosperma differs from P. virginica in having acuminate to mucronate (vs. obtuse) abaxial sepals, and broad (vs. narrow) seeds with shal- lowly (vs. deeply) concave adaxial sides. The Old World sect. LANcIFOLIA Barn. (sect. Arnoglossum Dene.) (oblan- ceolate leaves, grooved peduncles, dense spikes, and abaxial sepals fused for more than half their length) consists of perhaps six species, one of which is represented in our area. Plantago lanceolata, 2n = 12, 13, 24, 96, native to Europe and northern and central Asia, is established throughout most of the world, especially in temperate regions. It is an extremely variable species that is widely distributed across temperate North America (except the prairie prov- inces of Canada) and Mexico in most kinds of disturbed habitats. Perhaps the best-known member of the genus, P. /anceolata has been an extremely popular subject for research, some of which is summarized below. (See also Cavers et al. for a synthesis of much of the recent information concerning many aspects of its biology.) Plantago sect. GNAPHALOIDES Barn. (flowers paired or in threes; hairs of the scape wider than those of the leaves, and at least some hairs ascending) com- prises the American species included by Pilger in sect. LEucopsyLLium and was divided into four series by Rahn (1978a). The seven North American species, all 2n = 20, are in series Gnaphaloides Rahn (annuals with well-de- veloped taproots; linear leaves [but wider than those of species in sect. MICROPSYLLIUM]; spikes dense, cylindrical; bracts, especially of the lowermost flowers, often with an elongated green nerve; corolla lobes relatively conspic- uous and recurved, with cordate to obtuse bases; and anthers small). Five species, including the three that occur in our range, have zygomorphic corollas with cordate lobes (vs. actinomorphic with slightly cordate or obtuse lobes in the two other species). Of our species, P. aristata Michaux and P. Wrightiana Dene. (P. Hookeriana Fischer & Meyer var. nuda (Gray) Poe) are more com- mon and have adaxial leaf surfaces that are glabrous or nearly so; in the less common P. patagonica Jacq. (P. Purshii Roemer & Schultes) these surfaces are densely hairy. Plantago aristata is easily recognized by its conspicuous, often leaflike bracts that are usually smaller toward the apex of the spike. Taxonomically insignif- icant depauperate individuals with very narrow leaves, reduced flowers, and hairy, few-flowered spikes can be found, but such inflorescences occur on oth- erwise normal plants, and all intermediate forms exist. Plantago aristata 1s a common weed throughout the eastern United States (with the exception of the extreme north). It grows most commonly in dry, sandy soil along roadsides and in pastures, fields, and other kinds of recently disturbed habitats. It is an adventive in Oregon and California and in various parts of the Old World (Rahn, 1978b). Plantago Wrightiana differs most obviously from P. aristata in having shorter (and therefore less conspicuous) bracts. According to Rahn (1978b), the species occurs in three disjunct areas: central Arizona; Oklahoma, Texas, and adjacent parts of Mexico; and parts of Mississippi, Alabama, Geor- 1984] ROSATTI, PLANTAGINACEAE 547 gia, Florida, and the Carolinas. (I have, however, seen herbarium specimens from Louisiana and southwestern Arkansas that cast doubt on Rahn’s conten- tion that the southeastern populations are disjunct from those in Texas and Oklahoma.) Rahn also stated that, on the basis of ecological data, P. Wrightiana is probably not native to our area. The third of our species in series Gnaphaloides, Plantago patagonica, 1s intermediate in bract width and length between P. aristata and P. Wrightiana. It is at least an adventive in the Southeast. I have seen three specimens of P. patagonica (two were annotated by Rahn in 1975): one from railroad tracks in Cleveland Co., Arkansas, and two specimens from a roadside in Lancaster Co., North Carolina. (Smith has also reported it from Arkansas.) The species is otherwise known from southern and central Argentina; from an area between northern Baja California and south-central British Columbia eastward to the western Great Lakes and central Texas; and from two localities in interior New England (Rahn, 1978b). Species of sect. PsyLLium (Miller) Lam. & DC. (branched, caulescent herbs or subshrubs with opposite or rarely whorled leaves) are native to the Medi- terranean region and the Middle East. Plantago Psyllium L.,> 2n = 12, an annual with dense, subcapitate spikes and a taproot, has been reported from various parts of the world. In North America it has been collected in sandy and/or waste places. Plantago Psyllium is rare in southern Canada (Bassett, t commercial bird-seed (Munz), and well established in the eastern United States (Gleason & Cronquist). It is rare and perhaps not persistent in our area. Al- though it has been reported from coastal dunes in North Carolina (New Han- over Co.; Radford et a/.), I have seen neither a voucher specimen for this station nor any other specimen from the Southeast. Cytological studies have provided data that bear, either alone or in combi- nation with palynological information, on various systematic problems in the arisen b polyploidy, reduction in the number of chromosomes, and hybridization —the latter two processes often followed by chromosomal duplication), and that differences in chromosome morphology do not correlate with the two subgenera based on leaf arrangement (cf. Harms in Harms & Reiche; Pilger). The Old World and American species included by Pilger in sect. LeucopsyLLIUM were 3 There has been some confusion about the nomenclature of this species. Linnaeus (Species Plan- foliis mis, ade aphyllis” for P. Psyllium and ‘“‘caule ramoso herbaceo, fol. integerrimis, spicis ee for a new species, P. indica. Panigrahi has treated P. seers L. aes as an anes synonym of P Piha (1753), and th e P. aren Waldst. & Kit. has been used by Shear and by Chater and Cartier (in Moore, an ae listed (in synonymy) P. Psyllium as a nomen ambiguum and P. indica as a nomen illegitimum 548 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 considered by Rahn (1978a) to form morphologically distinct groups and were therefore placed by him in sections ALBICANS Barn. and GNAPHALOIDES, re- spectively. This position was reinforced by Bassett and Crompton, who ob- served that P. fastigiata E. L. Morris (P. insularis Eastw.) differs from the other North American species of sect. LEUCOPSYLLIUM (i.e., those included in sect. GNAPHALOIDES by Rahn) in several ways, including chromosome number 2n = 8 vs. 20) and pollen morphology (for example, smaller grains with operculate pores). Thus, although sections ALBICANS and GNAPHALOIDES appear to be taxonomically distinct, the former is not confined to the Old World and is therefore not geographically isolated from the latter, as was implied by Rahn (1978b). Finally, although both Wodehouse and Bassett (1973) have remarked that P. major and P. Rugelii differ more palynologically than morphologically, such differences could be due to chromosome number alone: the grains of P. Rugelii, a tetraploid, are larger and have more numerous pores than those of P. major, primarily a diploid. An interesting anatomical feature of Plantago is the occurrence of cambiform tissue in the cortex and/or pith of the fleshy rhizomes of many species, par- ticularly those in sect. PLANTAGO (Metcalfe & Chalk). These rows of cells appear to represent incompletely developed vascular bundles and are apparently con- nected with one another (but not with the bundles of the vascular ring). In P. major and P. media L. they become converted into true medullary vascular bundles. The tendency for insular members of a group to be more woody than their continental relatives is exemplified by species of Plantago (Carlquist). The ability to accumulate secondary xylem has evolved independently in this genus several times, apparently in response to the relative uniformity of island cli- mates. The adaptation of these same species to wetter conditions following migration from drier continental areas has involved an increase in vessel ele- ment length and diameter. In Plantago, as in other groups of predominantly herbaceous dicots, wood produced by insular species is largely rayless. Nodal anatomy in Plantago is extremely variable and a poor indicator of infrageneric relationships (Misra, 1966). Species in such dissimilar groups as sections CORONOPUS (Miller) Lam. & DC. (of subg. Coronopus (Miller) Rahn) and PsyLuium (of subg. PsyLiium) are unilacunar, while P. ovata Forskal (sect. ALBICANS of subg. PsyLLrum) is trilacunar and P. Lagopus L. (sect. LANCIFOLIA of subg. PsyLtium) and P. major (sect. PLANTAGO) are multilacunar. Anatom- ical complexity at the nodes appears to be a function of leaf size because unilacunar nodes are found in species with narrow leaves and bases and mul- tilacunar types are associated with broad leaves and sheathing bases. The presence of an endodermis with Casparian thickenings has been noted in both stems (Metcalfe & Chalk) and leaves (Trapp) in species of Plantago. Trapp found that in each of eight examined species (including P. lanceolata and P. major), the leaf endodermis is structurally no different than that found in angiosperm roots, and experiments showed that it is even potentially capable of arresting the outward diffusion of solutes. He considered it improbable, however, that such a function is actually realized and concluded that the pres- 1984] ROSATTI, PLANTAGINACEAE 549 ence of a foliar endodermis in these plants appears to be neither beneficial nor detrimental to a leaf’s functioning. Leaves of Plantago often resemble those of monocotyledons in their nar- rowness, parallel venation, and sheathing bases. Even species with distinct petioles and blades are often said to be pseudo-laminate, the blade representing (as in monocots) an “‘expansion of the apical region of the original phyllode” (Arber, 1918). Furthermore, leaf development involves the cessation of inter- calary activity and the subsequent maturation of tissues in a basipetal direction, as is well expressed in monocots but is also seen in conifers and other dicots with narrow leaves (Esau). The stomata of young leaves of Plantago are anomocytic (ranunculaceous); those of adult leaves are various (Moncontié). Stomata are usually diacytic (caryophyllaceous) in a number of species, including P. /anceolata and P. Psyl- lium, but they are rarely so in P. major. They may be distributed either on both surfaces of the leaf (e.g., as in P. lanceolata, P. major, and P. Psyllium) or (as in two extraregional species) only abaxially (Metcalfe & Chalk). In P. lanceolata and the extraregional P. altissima L. stomatal size appears to be positively correlated with chromosome number (Moncontié). The floral anatomy of Plantago lanceolata, P. major, and P. Rugelii has been studied by Henderson, whose observations suggested that the bract in Plantago actually represents a fifth sepal. Henderson observed, among nu- merous other things, that the stamens separate from the receptacle as a ring of tissue (remaining connate for some distance up the ovary) and are never fused to the corolla in P. lanceolata and P. major; my own observations, however, reveal that they are indeed epipetalous, as she correctly indicated for P. Rugelii. The ovule in Plantago has a single, massive integument and is tenuinucellar (the archesporial cell develops directly into the megaspore mother cell). It has been described as anatropous by Cooper (for P. /anceolata, although his illus- tration showed it to be more similar to what is usually considered hemianat- ropous), anacampylotropous by Misra (1964, for Plantago), hemianatropous to anatropous by Davis (for the family), anatropous to hemitropous by Cron- quist (1981, for the family), and more or less anatropous, erect, or anacam- pylotropous by Corner (for the family). Ovule and seed development in P. lanceolata have been described in great detail by Cooper (see also Davis; Misra, 1964), who stated that the megagametophyte in this species is of the Polygonum type. Although Cooper also indicated that the basal cell of the zygote forms the suspensor, Davis stated that the embryogeny is of the Onagrad type. The presence of both micropylar and chalazal haustoria in the developing seed was noted above (family discussion). Circumscissile dehiscence of the fruit occurs in such disparate genera as Plantago and Celosia (Amaranthaceae), Anagallis and Centunculus (= Ana- gallis) (Primulaceae), Portulaca and Lewisia (Portulacaceae), and Hyoscyamus (Solanaceae). In each of the species of Plantago studied by Rethke (including P. major and P. pusilla), the cells near the region of future dehiscence remain meristematic through much of fruit maturation. These cells are smaller and have thinner walls than adjacent epidermal and/or subepidermal cells. Dehis- 550 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 cence results from the breaking and separation of cells in this zone of mechanical weakness and occurs as the drying capsule shrinks against the mature, unyield- ing seeds (see also Lamba & Gupta). The extent to which genotype (vs. environmental factors) determines phe- notype, including morphological oddities, has received considerable attention. Evidence has suggested that while branched spikes are under genetic control in P. lanceolata (Weatherwax; cf., however, Yamaura) and P. major (Ham- marlund), they seem to result from external factors in P. Rugelii (Weatherwax). The lengths of scapes, capsules, seeds, and bracts (as well as the number of flowers produced) were all found to be positively correlated with plant size in P. aristata (Goodwin), which is evidently not genetically determined. It is notable that depauperate plants sometimes produce lowermost bracts that do not exceed the flowers they subtend, and that they may thus rather closely resemble several other species of sect. GNAPHALOIDES. In a series of long-range experiments involving both cuttings and seedlings, Marsden-Jones and Turrill found that plant size and peduncle pubescence in Plantago major were mostly determined by the soil in which the plants were grown (cf., however, Warwick & Briggs, 1979). A number of studies have been undertaken to determine the relationship between phenotype and ploidy level in Plantago. Chandler (1954) showed that pollen size, seed size, quantity of mucilage, and plant vigor are all greater in colchicine-induced tetraploids (compared to the corresponding diploids) in P. rhodosperma, P. Wrightiana, and the extraregional P. ovata. In P. ovata such treatment resulted in larger leaves with generally larger cells, larger stomata, and broader epidermal hairs (Chandler & Barton). Several physiological dif- ferences were also noted, including the fact that tetraploids are, in general, developmentally slower than diploids. It is also interesting that the amount of aucubin (see family discussion) did not differ between the two groups. In other studies involving extraregional species, Rahn (1954) discovered that in P. media the seed number per capsule, scape length, and leaf size are all greater in natural tetraploids than in diploids (although pollen diameter and stomatal size do not differ), and Bocher et a/. found in P. Coronopus L. that the anthers, flowers, and plants themselves are larger in natural hexaploids than in diploids. Hybrids involving species of Plantago represented in the Southeast have apparently not been encountered (with the exception of the artificially produced P. cordata x P. eriopoda [Tessene, 1969]). Artificial crosses between P. major (sect. PLANTAGO) and P. /anceolata (sect. LANCIFOLIA) were unsuccessful (Rahn, 1957). In fact, hybrids involving these two species, either together or with other species, are not known (Sagar & Harper, 1964). No hybrid in which P. major was a parent has been reported, and the identity of herbarium specimens of a putative hybrid between P. /anceolata and the extraregional P. media (sect. LAMPROSANTHA Dene. of subg. PLANTAGO) is doubtful. All attempts to produce artificial hybrids between P. media and either P. major or P. lanceolata have failed. Attempted crosses between extraregional species in sect. LAMPROSANTHA and those in sects. PLANTAGO, LANCIFOLIA, and Coronopus have also been unsuccessful (Rahn, 1957). Also among species not occurring in the Southeast, natural introgressive hybridization was reported in subg. CORONOPUS sect. 1984] ROSATTI, PLANTAGINACEAE 551 Maritima H. Dietr. (Cartier, 1971, 1972), and a high degree of interfertility was observed in subg. PsyLtium sect. MONTANA Barn. when artificial crosses involved plants with the same chromosome number (Cartier, 1973). The occurrence of perfect and carpellate flowers on separate plants (gyno- dioecy) has been observed, investigated, and discussed since the time of Darwin. Plantago lanceolata, in particular, has been the subject of a number of inves- tigations (see Bartlett, Ludwig, and Ross [1969]). Ross proposed that there are dominant alleles at two loci for perfect flowers, so double homozygous recessive individuals produce only carpellate flowers (cf., however, Atal, Van Damme & Van Delden, and Krohne et al. regarding a possible cytoplasmic effect). Because such individuals contribute genes to subsequent generations through ovules only, it is reasonable to postulate the existence of a compensatory mechanism by which carpellate-flowered plants are maintained in populations in which plants with perfect flowers contribute genes through both ovules and pollen and are therefore presumably at a selective advantage. Krohne et al. determined, in fact, that in natural populations (in northern California) car- pellate plants generally produce more seeds than do those with perfect flowers (see also Primack, 1978). Plantago lanceolata is also self-incompatible, even though such a system might appear to reinforce the process of outcrossing unnecessarily. Investiga- tions by Ross (1973) suggested that at least four alleles at a single locus are involved, and that incompatibility results if the pollen has either of the alleles present in the style. (Unlike most other angiosperms with trinucleate pollen, the incompatibility is gametophytic.) Ross (1973) argued that matings between wholly perfect or wholly carpellate progeny (which could result from individual maternal parent plants with perfect or carpellate flowers, respectively) would be limited by self-incompatibility in the first case (there would be no limitation on such crosses under gynodioecy alone) and prevented by gynodioecy in the second (a portion of such crosses would be possible under the incompatibility system alone). In this way the two systems would be mutually reinforcing. Plantago is usually considered to be primarily wind pollinated, at least in those species with chasmogamous flowers, and a majority of the evidence seems to bear this out. Many of the traits usually associated with anemophily (flowers proterogynous, small, unattractive, scentless; stigmas large, pubescent; anthers large, versatile on long filaments; and pollen spheroidal, dry, relatively smooth, with little variation in size) are found in species of P/antago. For example, on the basis of data gathered by Bassett and Crompton from North American plants, Primack (1978) demonstrated that variation in mean pollen diameter was greater among species with cleistogamous flowers than among those with chasmogamous ones. This presumably reflects selection against pollen that is either too large or too small (see also Proctor & Yeo Some species of Plantago appear to be at least pattly entomophilous (but, as Proctor & Yeo have pointed out, bees sometimes collect pollen from plants that are anemophilous). Clifford stated that in Britain P. /anceolata is consid- ered to be wind pollinated, but plants of it are regularly worked by honey bees for pollen whenever conditions are warm enough and a sufficient number of spikes are flowering within a few feet of one another. A more elaborate study Doz JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 of insect pollination in this species in the Netherlands (Stelleman & Meeuse) attempted to prove that pollen is actually transferred to the stigmas by flies of the genera Melanostoma and Platycheirus (Diptera: Syrphidae). Spikes of a group of donor plants were dusted with stained pollen of P. /anceolata. This pollen was later detected on both the flies and the stigmas of receptor plants located in a separate group upwind, while only negligible amounts of it were found on the donor plants’ side of test slides placed between the two groups of plants. Several investigations of reproductive biology have involved species of P/an- tago in the Southeast. Primack (1979) estimated that “reproductive output” is greater among annuals (vs. perennials), spring (vs. summer) annuals, and weedy (vs. nonweedy) perennials. Primack and Antonovics (1982) later found that in “reproductive effort” eight populations of P. /anceolata from the central Pied- mont of North Carolina had a range of values comparable to that obtained by Primack (1979) for various perennial and/or weedy species of Plantago, but less than that determined for annual species (although the degree of genetic control was not clearly established). Primack (1980) also determined that phe- notypic variation in Plantago, as measured by several criteria (number of inflorescences, number of capsules per inflorescence, number of seeds per cap- sule, and weight per seed), is no less in rare species than in more widespread ones. Significant differences in all of these measures except seed number per capsule were later observed under controlled conditions in P. /anceolata (Pri- mack & Antonovics, 1981). Studies involving Plantago lanceolata have shown that these plants may become physiologically adapted to very local conditions. Among populations in the Southeast, Wu and Antonovics (1976) demonstrated a greater tolerance to lead by roadside (vs. nonroadside) individuals, and Pollard showed essen- tially the same kind of response to zinc, copper, arsenic, and lead at sites contaminated with these metals. In contrast to various authors who have sug- gested that the wide geographic ranges characteristic of many weedy species are primarily the result of broad temperature optima for photosynthesis, Tera- mura and Strain demonstrated that substantial differences exist in the photo- synthetic response to temperature among populations growing in relatively close proximity (eight meters to ten kilometers apart). Plants cloned from leaves taken from a shaded population were more sensitive to temperature extremes than were those propagated from open and sun-flecked populations that were exposed to much greater diurnal and seasonal fluctuations in temperature. Plants from the open population had much greater light-saturated rates of photosynthesis as well. Plantago pollen has been helpful in studies regarding the effects of human activity on vegetation. Godwin (1975) pointed out that the increase in the number of records of P. major and P. lanceolata in British postglacial sediments corresponds to the praetess of deforestation and the spread of scum by Neolithic man. (There a weedy species as P. media and P. maritima Ex) Tn the Vosges region of aaa the degree of aforestation and cultivation is reflected in the relative abundance of the pollen of P. /anceolata in surface samples (Janssen). 1984] ROSATTI, PLANTAGINACEAE 553 The rather substantial, but frequently underestimated, economic significance of Plantago has been thoroughly discussed by Hammond. Several species, but especially P. lanceolata and P. major, are well known and noxious lawn weeds that also compete with crop plants for light, space, and nutrients (see Ham- mond; Higbee & Lee). Inadvertent dissemination of these plants results from Plantago seeds being unavoidably harvested with (and difficult to separate from) crop seeds. The leaves of Plantago do not dry readily and can therefore contribute to spoilage in hay that is sufficiently contaminated (Gill & Vear). Plantago species also act as hosts to a variety of viruses and (other) organisms known to cause disease in crops including beets, potatoes, turnips, eUcUMDETS; tomatoes, broad beans, and tobacco. Precisely how tl loss is largely a matter of speculation at present. Plantago lanceolata is sometimes fed to livestock as a mineral supplement, and it is still grown in the Netherlands for use in herbal remedies. Plantago major has also been used in this way for a variety of medicinal problems. The mucilaginous seeds of P. Psyllium and related species (“‘psyllium”’ seeds) act as a mild laxative and are harvested from crops raised specifically for this purpose. Chandler (1954) reported that almost four million pounds of psyllium seeds were imported into this country in 1953 and discussed the possibilities of domestic production for use of the mucilage in pharmaceutical preparations and as basic stabilizers in making ice cream, in printing and finishing, and in setting lotions (see also Morton). According to the United States Department of Agriculture, Foreign Agricultural Service, over 12 million pounds of psyllium seeds were imported into > this country in 1981, at a value of over nine million 1981 dollars. REFERENCES: Q 1975, 1980), GLEASON & CRONQUIST, GODWIN (1 975), Harms & REICHE, MCCULLAGH, Moncontié, Moore (1976), and PILGER Autes, H. E., C. R. Bett, & A. E. RADForD. Species new to the flora of North or South rolina. Rhodora 60: 10-32. 1958. [P. Hookeriana var. nuda (Gray) Poe (= P Wrightiana) established in the Carolinas.] Antonovics, J., & R. B. Primack. Experimental ecological genetics in Plantago VI. The pee of seedling vans of P. lanceolata. Jour. Ecol. 70: 55-75. han genetics in determining mortality, growth rate, and fecundity ArsBerR, A. The aisilede theory of the monocotyledonous leaf, with special reference to anatomical evidence. Ann. Bot. 32: 465-501. 1918. ATAL, C. K. Cytoplasmic male sterility in psyllium (Plantago ovata Forsk.). Curr. Sci Bangalore 27: 268. 1958. [Crossing experiments gsi cytoplasmic franemission: see also VAN DAMME & VAN DELDEN and Kroune et al. Baker, H. G. Evolutionary mechanisms in ie aman bole. Science 139: 877-883. 1963. [P. lanceolata said to be gynodioecious, with as much as 70 percent of some populations consisting of plants with ay eae flowers. ] BaLpo, B. A., Q. J. CHENSEE, M. E. H. Howpen, & P. J. Sahar. “ilgieens from plantain 554 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 (Plantago lanceolata)[;] studies with pollen and plant extracts. Int. Arch. Allergy Appl. Immunol. 68: 295-304. 1982. [At least six of 16 pollen antigens allergenic; allergenic role for dicot plant fragments (other than pollen) suggested for the first time. ] BarTLETT, H. H. On gynodioecism in Plantago lanceolata. Rhodora 13: 199-206. 1911. ee flower forms. ance of sex forms in Plantago lanceolata. Ibid. 15: 173-178. 1913. [In- eee flower forms. ] Bassett, I. J. Taxonomy of North American Plantago L., section Micropsyllium Decne. Canad. a - 44: 467-479. 1966. Tax y of Plantago L. in North America: pte Holopsyllium Pilger, Palacopsin Pilger, and Lamprosantha Decne. Ibid. 45: 565-577. 1967. plantains of Canada. 47 pp. Research Branch, Canada Dep. Agr. Ottawa. 1973. includes keys, synonymy, descriptions, comments, illustrations, Sabin maps; treats eight spp. reported from the Southeast.] . R. BAum. Conspecificity of Plantago fastigiata of North America with P. ovata of the Old World. Canad. Jour. Bot. 47: 1865-1868. 1969. [Plants assigned to P. fastigiata are in fact the Old World P. ovata, which became established during the late 18th and early 19th centuries in California: see also BAsseTT & CROMPTON and Stespins & Day BERENDSE, F. Competition between plant populations with different rooting depths III. Field experiments. Oecologia 53: 50-55. 1982. [Utilization of nutrients from deeper soil layers by P. lanceolata during interspecific competition BLom, C. W. P.M. Effects of trampling and soil compaction on the occurrence of some Plantago species in coastal sand dunes II. Trampling and seedling establishment. Oecol. Pl. 12: 363-381. 1977. B6cHerR, T. W. Studies on variation and biology in a lanceolata L. Dansk Bot. Ark. 11(3): 1-18. 1943. [Genotype, age of plant, and environment interacted in cultivation to determine leaf length and fe scape length, and spike length and number. , K. Larsen, & K. RAHN. Experimental and cytological studies on plant a I. Kohlrauschia prolifera and Plantago Coronopus. Hereditas 39: 289-304. Brown, M. J. An experimental taxonomic study of Plantago tasmanica Hoo . f. ae P. Daltonii Dene. Austral. Jour. Bot. 29: 441-452. 1981. [Common-garden exper- iments; introgression. ] BunTING, A. H. Some reflections on the ecology of weeds. Pp. 11-26 in J. L. HARPER, ed., The biology of weeds. Oxford. 1960. [“‘Late glacial” listed as time of apparent entry of P. lanceolata, P. major, and P. media into the British flora. CARLQuIST, S. Wood anatomy of insular species of Plantago and the problem of ray- lessness. Bull. Torrey Bot. Club 97: 353-361. 1970. Cartier, D. Etude biosystématique de quelques espéces du genre Plantago (Tourn.) L. (sections Coronopus D. C. et Oreades Decne.). I.—Historique, races chromoso- miques du Plantago alpina L. et du oe serpentina All. (English summary.) Revue Gén. Bot. 78: 493-556. 1971. I.—La variation au sein de différentes po- pulations naturelles ou re en : Plantago alpina L. et du Plantago ser- pentina All. (English summary.) /bid. 79: 201-248. 1972. II].—Apports de croise- ments expérimentaux chez les P. alpina L. et P. serpentina All. Conclusions sur la complexe P. alpina-P. serpentina. (English summary.) [bid. 80: 5-40. 1973. IV La section Oreades Decne. Sea summary.) Ibid. 129-149, Cavers, P. B., I. J. Bassett, & C. W. Crompton. The biology of Canadian weeds. 47. erie lanceolata L. Canad. tour Pl. Sci. 60: 1269-1282. 1980. [Morphological riation, economic importance, geography, habitats, history (first reported in Nova aa in 1829), growth and development, reproduction, hybridization, population 1984] ROSATTI, PLANTAGINACEAE 55) dynamics, and response to human manipulations (including herbicides and other chemicals) and parasites.] aara: C. Seed germination for some species of needs Contr. Boyce Thompson 7: 265-271. 1953. [P. rhodosperma, P. Wrightia mprovement of Plantago for mucilage Sa eace and growth in the United States. Ibid. 495-505. 1954. [P. rhodosperma, P. Wrightian & . Barton. Morphological and physiological aa of diploid and tet- raploid ‘Plantago ovata Forsk. Contr. Boyce Thompson Inst. 18: 193-214. 1955. CurForD, H. T. In oe pollination of Plantago lanceolata L. Nature 193: 196. 1962. Cute, W. N. The meaning of plant names LXIII[.] The plantains. Am. Bot. 41: 103- we 1935. [itor alae CoLeMAN, R. W., & S ELSH. Plants from northern Manitoba, Canada and their meee (Abstr.) re Iowa Acad. Sci. 88(Abstracts): 13. 1981. [P. major in calcareous soils (pH ca. 8). Cooper, G. O. Be iconen of the ovule ee i formation of the seed in Plantago lanceolata. Am. Jour. Bot. 29: 577-581. CorreLL, H. B. Plantaginaceae. Pp. 1475- rae in 1D. S. CorRELL & M. C. JOHNSTON. Manual of the vascular plants of Texas. Renner, Texas. 1970. [12 spp.; P. hetero- phylla Nutt. indicated as a questionable synonym of P. hybrida W. Barton, see GopFREY & WOOTEN Crompton, C. W. Pollen grains and biosystematics. Canad. Jour. Bot. 60: 294-300. l CUNNINGHAM, A. M. Certain chemical features in the seeds of Plantago Haat and P. patagonica. Proc. Indiana Acad. Sci. 1894: 121-123. 1895. [A blue pigment, thought to be a glucoside related to indican, released upon soaking in ihe A revision of the species of the genus Plantago occurring within the United States. Ibid. 1896: 190-207. 1897. [17 spp., eight credited to the Southeast; P. pusilla Nutt. a synonym of P. elongata Pursh; P. major “from Delaware to California and Damme, J. M. M. van, & W. vAN DELDEN. Gynodioecy in Plantago lanceolata L{.] I. Polymorphism for plasmon type. Heredity 49: 303-318. 1982. [Two series (from sterile to fertile) of stamen forms, one showing size reduction and the other involving petaloid stamens; sala of gynodioecy both Se ae and nuclear, see also ATAL and KRroune et al.; see also Heredity 52: 77-93. D’Arcy, W. G. Plantagnacee. In: R. E. Woopson, Jr., & a ~ ‘SCHER y, eds., Panama. Ann. Missouri Bot. Gard. 58: 363-369. 1971. [Three spp.; P. /anceo oe ta not reported from below 3300 ft, P. major from as low as 200 ft; neither considered mmon enough to be a nuisance.] eee H. Studien an cubanischen Plantaginaceae. Wiss. Zeitschr. Friedrich Schiller Univ. Jena Math. Nat. 28: 701-708. 1979. [P. lanceolata, P. major, P. Rugelii at or near sea level; floral morphology, cytology, palynology; illustration, map of dis- tribution in Cuba.] Dopps, J. G. Plantago Coronopus L. Jour. Ecol. 41: 467-478. 1953. (Systematics and biology; cementing of seed to substrate by mucilage thought to be advantageous in germination Dow Lina, R. E. “The structure of the ovary in the genus Plantago L. — I. The British species. Jour. Linn. Soc. Bot. 50: 323-336. 1936. [P. lanceolata, P. major, variation in seed number in P. major suggests the ovary is of little value in defining either sections or species.] Eames, A. J. Morphology of the angiosperms. Frontisp. + xiii + 518 pp. New York, Toronto, and London. 1961. [Anemophily in Plantago; genus provides example of reduction to one flower from racemose clusters; leaves phyllodes.] Esau, K. Plant anatomy. xii + 735 pp. New York and London. 1953. [Plantago, 421 556 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 (striate ema 455 (leaf development), 581 (circumscissile dehiscence), 604 (mu- cilaginou Foster, R. C. The publication-date of Nuttall’s “Arkansas Flora.”” Rhodora 46: 156, 157. 1944. [Protologue of P. heterophylla published in 1835.] Fow er, N. L., & J. ANronovics. Smale-scale variability in the demography of trans- plants of two herbaceous species. Ecology 62: 1450-1457. 1981. [P. lanceolata; high summer temperatures and consequent water shortages seemed to pose greater risks than did low winter temperatures and freezes. ] Fusyrwara, I. Karyotype analysis in ped I. (In Japanese; English summary.) Kro- — 27/28: 962-968. 1956. otype Srraee 2 oe IL (In Japanese; English summary.) Jap. Jour. Genet 31: 184-191. 6. [P. lanceolata, P. Psyllium, P. virginica, karyotypes of ubg. Psyllium een similar to those of subg. Plantago; P. virginica an allo- eed P. Psyllium and P. indica “partially related” karyotypically.] GILL, N. T., & K. C. VEAR. eee botany. viii + 636 pp. London. 1958. [Plan- taginaceae, 451; seedlings of P. major and P. lanceolata illustrated, 496, 497.] GoprFrey, R. K. Plantago cordata still grows in Georgia. Castanea 26: 119, 120. 1961. {Confirmed for Gordon Co.] & J. W. Wooten. Aquatic and wetland plants of southeastern United States. Dicotyledons. x + 933 pp. Athens, Georgia. 1981. [Plantago, six spp., 710-712.] Gopwin, H. Age and origin of the ‘Breckland’ heaths of East Anglia. Nature 154: 6, 7. 1944. [Pollen of P. lanceolata indicates origin in Neolithic forest clearance.] Goopwin, R. H. Variation in a natural population of Plantago aristata. Bull. Torrey Bot. Club 76: 109-115. 1949. GorENFLOT, R. Introgression, polymorphisme et taxonomie chez les Plantaginacées. Adansonia, II. 4: 393-417. 1964. [Natural and artificial hybridization between and within P. Coronopus and P. macrorhiza Poiret. Grecor, J. W. Experimental taxonomy IV. Population differentiation in North Amer- ican and European sea plantains allied to Plantago maritima L. New Phytol. 38: 293-322. 1939. [Spike density, sepal length/width, growth habit, scape length follow an ecological gradient; leaf pubescence varies geographically from western North America to central Europe.] GrirFiTHs, B. M. Growth experiments on Spergula and Plantago. Jour. Bot. London 60: 228-230. 1922. [P. lanceolata var. sphaerostachya Roehl. an environmental modification; a form of P. major with large inflorescences (and other characters) suspected of being under control of a recessive allele. HamI_ton, K. C., & K. P. BucHHOLTz. Effect of rhizomes of quackgrass (Agropyron repens) and shading on the seedling development of weedy species. Ecology 36: 304— 308. 1955. [Effect positive on P. Rugelii.] HAMMARLUND, C. Uber die Vererbung anormaler Ahren bei Plantago major. (English summary.) Hereditas 2: 113-142. 1921. [Alleles for branched spikes with leaves instead of bracts considered recessive. ] Hamner, K. Entwicklungstendenzen bliitendkologischer Merkmale bei Plantago. (En- glish summary.) Flora 167: 41-55. 1978. [Entomophily considered primitive within ] Plantago. oe J. Plantago as a host of economically important viruses. Advances Virus 7: 103-140. 1982. aie A. Plantaginaceernes og Lentibulariaceernes udbredelse i 1 Danmark. (English summary.) Bot. Tidsskr. 56: 1-35. 1960. [P. Rug northern Europe. ] Hareer, J. L., J. T. Wittiams, & G. R. SAGAR. The behavior of ne in soil I. The heterogeneity of soil surfaces and its role in determining the establishment of plants from seed. Jour. Ecol. 53: 273-286. 1965. [Experiments involving P. lanceolata and , |] Harper, R. M. Notes on Plantago, with special reference to P. cordata. Castanea 9: 1984] ROSATTI, PLANTAGINACEAE PY, 121-130. 1944. [P. elongata, P. aA and P. pusilla; historical notes on the occurrence of P. cordata in the South HawTuorn, W. R., & P. B. CAvErRs. eas on dynamics of the perennial herbs P/an- tago major L. and P. Rugelii Decne. Jour. Ecol. 64: 511-527. 1976. & Dry weight and resource allocation patterns among individuals in populations of Plantago major and P. Rugelii. Canad. Jour. Bot. 60: 2424-2439, 1982. [Greater overwintering survival and longevity observed in natural populations of P. Rugelii.] HENDERSON, L. B. Floral anatomy of several species of Plantago. Am. Jour. Bot. 13: 397-405. 1926. Hiasee, E. C., & A. Lee. Drug and medicinal crops. Pp. 127-160 in C. M. Wizson, ed., New crops for the New World. New York. 1945. [Psyllium, 148, 149; sea eee fertilization when grown commercially.] HILLEBRAND, W. Flora of the Hawaiian Islands: a description of their phanerogams and vascular cryptogams. Frontisp. + xcvi + 673 pp. + 4 maps (incl. 3 foldouts). Lon- don, New York, and Heidelberg. 1818. [P. major introduced and said to have 6-8 inch petioles and 1.5-2 foot spikes on the high pasture lands; two native species endemic; cf. oe Hitcucock, C. L., & A. Cronquist. Flora of the Pacific Northwest. xix + 730 pp. Seattle and London. 1973. [Plantaginaceae, 447, 448] Hype, B. B. Differentiated chromosomes in Plantago ovata. Am. Jour. Bot. 40: 809- 815. 1953. [Chiasma frequency, chromosome differentiation, heterochromatic seg- ments.] Mucilage-producing cells in the seed coat of Plantago ovata: developmental fine structure. Ibid. 57: 1197-1206. 1970. [Anatomical, ultrastructural, and histochem- Hype, H. A., & D. A. WittiaMs. Studies in atmospheric pollen III. Pollen production and pollen incidence in ribwort plantain (Plantago lanceolata L.). New Phytol. 45: 271-277. 1946. [Low delay or suppress daily ee time; estimated that almost 94 percent of : all pollen liberated becomes airborn JANSSEN, C. R. Contemporary pollen assemblages from the Vous (France). Rev. Pa- laeobot. Palynol. 33: 183-313. 1981. JOHNSON, M. F. Phrymaceae and Plantaginaceae in Virginia. Digare Jour. Sci. II. 32: 12-16. 1981. [P. Psyllium from four counties; cf. RADFo al.] Karey, 8. L. M. Plantago major L.—spray aia cao Watsonia 9: 44, 1972. bas as inflorescences, funnel-leav KNosLocu, I. W. Intergeneric hybridization in ewe plants. see 21: 97-103. 1972, [Four intergeneric and interspecific hybrids in the Plantaginaceae.] Kroung, D. T., I. BAKER, & H. G. BAKER. The maintenance of the Be alieseats breeding system in Plantago lanceolata L. Am. Midl. Nat. 103: 269-279. 1980. [Poorly understood cytoplasmic inheritance factor” invoked to explain inconsist- encies in data; see also ATAL and VAN DAMME & VAN DELDEN.] KunkeEL, G. Uber T einige morphologische Anomalien bei Plantago lanceolata L. und anderen Pflanzen in Valdivia (Siidchile). Beitr. Biol. Pflanzen 34: 509-530. 1958. [Branched inflorescences and rosette formation on scapes.] Kuo, S. H., & W. H. Murpy. The response of city and rural populations to an urban environment. (Abstr.) ASB Bull. 20: 65. 1973. [Garden experiments with P. aristata.] LAMARCK, J. B., & A. P. bE CANDOLLE. Plantaginées. Fl. Franc. ed. 3. 3: 407-418. 1805. [Sections Plantago, Psyllium, and Coronopus established. Lapa, L. C., & V. Gupta. Anatomy of circumscissile dehiscence in Plantago ovata Forsk. Curr. Sci. Bangalore 50: 541-543. 1981. [Scanning electron micrographs of capsule.] Larsen, L. M., O. OLSEN, & H. SorENSEN. Failure to detect oe in Plantago species. eae pee 22: 2314, 2315. 1983. [Cf. RODMAN e. 558 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 LeBebev-Kosov, V. I. Flavonoids of Plantago major. Chem. Nat. Compds. 12: 730. 1977. [Translation of Khimiya Prirodnykh Soedinenii 6: 812, 813. 1976; evidently the first report of flavonoids in the Plantaginaceae (apigenin and two luteolins).] Lona, R. W., & O. Laketa. A flora of tropical Florida. xvii + 962 pp. Coral Gables, Florida. 1971. [P. aristata, P. lanceolata, P. major, and P. virginica Lupwic, F. Ueber die Blutenformen von Plantago lanceolata L. und die Erscheinung der Gynodiocie. Zeitschr. Naturw. 52: 441-449. 1879. [Intermediate flower forms. ] Mark, A. F. The flora of the grass balds and fields of the southern Appalachian Moun- tains. Castanea 24: 1-21. 1959. [P. cordata, P. lanceolata, P. major, and P. Rugelii.] Marsben-JOnes, E. M., & W. B. TurriLt. Transplant experiments of the British Eco- logical ee at Potterne, Wiltshire[.] Summary of results, 1928-37. Jour. Ecol. 26: 380-389. 1938. Martin, A. C. The comparative internal morphology of seeds. Am. Midl. Nat. 36: 513- 660. 1946. [Nonstarchy endosperm, embryo large and erect, cotyledons expanded in Plantago.] MEAGHER, T. R., J. ANTONOVICS, & R. PRIMACK. Experimental ecological genetics in Plantago. Il. Genetic variation and demography in relation to survival of Plantago cordata, a rare species. Biol. Conserv. 14: 243-257. me [Populations in North Carolina showed shifts in vigor as well as overall declin METCALFE, C. R., & L. CHALK. Plantaginaceae. Anat. Dicot, 2: 1053-1059. 1950. Misra, R. C. Development and structure of angiosperm seed—TIII[.] Plantago L. Bull. Natl. Bot. Gard. Lucknow 105: 1-14. 1964. . Morphological studies in Plantago II]. Nodal anatomy. Proc. Indian Acad. Sci. B. 63: 271-274. Morris, E. L. A revision 1 of the species of Plantago commonly referred to Plantago eee ae Bull. Torrey Bot. Club 27: 105-119. 1900. [P. patagonica said t to occur in North America; P. Purshii reported from Arkansas. ] North eee Plantaginaceae—III. Ibid. 36: 515-530. 1909. [P. pusilla com- pared with P. elongata, P. heterophylla, and P. hybrida;, cf. GopFrey & WoorTEN.] Morton, J. F. Major medicinal plants[.] Botany, culture, and uses. x1x + 431 pp. Spring- field, Illinois. 1977. [Plantago, 323- 332.} | — G.A.,&J.N. FInpLay. R d colonization in Canadia s. Canad. Jour. Bot. 48: 859, 860. 1970. ee flowers of P. major set viable ery (i.e., the plants are self-compatible and/or apomictic).] Munz, P. A. A California flora. vii + 1681 pp. Berkeley, Los Angeles, and London. 1959. [16 spp. of Plantago, 405- < also 78, 79 in Supplement (224 pp.), 1968.] Neva, A. C., & E. B. FiscHer. A comparative pharmacognostical study of Plantago Purshii R. & S. and Plantago ee Michx. seeds with the present N. F. VIII Plantago seeds. Jour. Am. Pharm. Assoc. Sci. Ed. 38: 34-41. 1949. [Detailed de- scriptions of seed structure. ] Nog, R., & C. W. P. M. BLom. Sir aates of three Plantago species in coastal dune grasslands i in relation to pore-volume and organic matter content of the soil. Jour. Appl. Ecol. 19: 177-182. 1981. [P. ee Pom Orum, G. Beitrag zur Kenntnis der Blattanatomie und cee von Plantago media - major L. und PI. lanceolata L., mit besonderer Berucksichtigung der Unter- scheidungsmiglichkeit der Blatter auch in Bruchstiicken. Beih. Bot. Centralbl. 50(Abt. cells were regular (six bivalents); cf., however, MCCULLAGH regarding chromosome morphology.] PaniGRAHI, G. A note on Plantago exigua (Plantaginaceae) and certain related taxa. Kew Bull. 30: 669-673. 1975 [1976]. Por, I. A revision of the Plantago patagonica group of the United States and Canada. Bull. Torrey Bot. Club 55: 406-420. 1928. [‘... plants growing in full sun tend to 1984] ROSATTI, PLANTAGINACEAE 559 have shorter, more vigorous stems and wider leaves, yet bracts [length and shape] remain fairly constant.’ > Jacquin’s description and plate of P. patagonica did not from Argentina were similar to P. Purshii. Type of P. Wrightiana considered to belong to P. Hookeriana var. nuda.] PoLLarD, A. J. Diversity of metal tolerances in Plantago lanceolata L. from the south- eastern United States. New Phytol. 86: 109-117. 1980. PRIMACK, R. B. Evolutionary aspects of wind pollination in the genus Plantago (Plan- taginaceae). New Phytol. 81: 449-458. 1978. [Carpellate plants produced more inflorescences and heavier seeds than did perfect ones; differences in seed weight attributed to the high cost of pollen production . Regulation of seed yield in Plantago. Jour. Ecol. 66: 835-847. 1978. [Includes data from all of the southeastern species. ] . Reproductive effort in annual and perennial species of Plantago (Plantagina- ceae). Am. Nat. 114: 51-62. 1979. [Includes data from natural populations and ee ae specimens representing all of the southeastern species. otypic variation of rare and widespread species of Plantago. Rhodora 82: 87- 95. 1980. [Includes data from all of the southeastern species.] & J. ANTONOViIcs. Experimental ecological genetics in Plantago. V. Components of seed yield in the ribwort plantain Plantago lanceolata L. Evolution 35: 1069- 1079. 1981. [In plants under controlled conditions, significant differences in inflo- rescence number per plant, capsule number per inflorescence, and seed weight, but ut in seed number per capsu & Experimental ecological genetics in Plantago. VI. Reproductive effort in populations of P. lanceolata L. Ibid. 36: 742-752. 1982. Proctor, M., & P. YEo. The pollination of flowers. 418 pp. + 56 black & white + 4 color pls. "New York. 2, RApForpD, A.E., H. E. AHLEs, & C. R. BELL. Manual of the vascular flora of the Carolinas. lxi + 1183 pp. Chapel Hi ll, Sas Carolina. 1968. Raun, K. Experimental and cytological studies in Plantago media. Bot. Tidsskr. 51: 300-307. 1954. Chromosome numbers in Plantago. Ibid. 53: 369-378. 1957. [Seeds referred to ‘the hybrid P. media x lanceolata always yielded one or the pgs of the parent species, unsuccessful Crosses attempted between P. /anceolata, P. major, and P apoE, two of 42 “‘strains’ of lanceolata, 2n= 12, reported as aneuploids, 2n= 13.] . Plantago section Virginica. Danek Bot. Ark. 30(2): 1-180. 1974. [28 spp.] . Plantaginaceae. Fl. Ecuador 4: 23-38. 1975. [Ten spp. of Plantago, Pee eae and P. major, none at low elevations.] menclatorial changes within the genus Plantago L., infraspecific taxa and ee of the genus. Bot. Tidsskr. 73: 106-111. 1978a. [Subg. Plantago, 11 sections, need of revision noted; subg. Coronopus established, two sections; subg. Psyllium, seven sectio Plantago ser. Cae Rhan[,] a taxonomic revision. [bid. 73: 137-154. y READ, D. J. H. K. Koucuek1, & J. HopGsSon. Vesicular-arbuscular mycorrhiza in natural vegetation systems I. The occurrence of infection. New Phytol. 77: 641-653. 1976. [Seedlings of P. lanceolata with mycorrhizal fungi.] RETHKE, R. V. The anatomy of circumscissile dehiscence. Am. Jour. Bot. 33: 677-683. 1946. [P. major, P. pusilla.) Rezk, M. R. Seed structure as a phylogenetic criterion. A case of Plantago seed. Egypt. Jour. Bot. 23: 51-62. 1980. [P. sees P. major, P. virginica. Rock, J. F. The genus Plantago in Hawaii. Am. Jour. Bot. 7: oe aa 1920. [P. lanceolata, P. major, and P. virginica ere tee | two native species e RODMAN, J. E. Divergence, convergence, and parallelism in Siete characters: 560 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 the glucosinolate-myrosinase system. Pp. 43-79 in D. A. Younc & D. S. SEIGLER, eds., Phytochemistry and angiosperm Phylogeny. New York, 1981. [Myrosin | cells (protein-rich idioblasts in which absent from the Plantaginaceae, even though hydrolysis products of glucosinolates Ross, M. D. Digenic inheritance of male sterility in Plantago lanceolata. Canad. Jour. Genet. Cytol. 11: 739-744. 1969. [Nearly continuous variation between plants with only perfect flowers and those with only carpellate ones.] . Breeding systems in Plantago. Heredity 25: 129- 133. 1970. [12 spp., including P. aristata and P. rhodosperma (assumed to be self-compatible because of cleisto- gamy), P. /anceolata (both gynodioecious and self-incompatible), P. major (both gynodioecious and self-compatible, at least locally), and P. Wrightiana (assumed to be self-compatible because fertilization prior to corolla expansion). . Inheritance of self-incompatibility in Plantago lanceolata. Ibid. 30: 169-176. RYMKIEWICZ, A. Studies on the species of the genus Plantago L. with reference to no vated for four years; illustrations of seeds and of transverse sections of endosperm and testa; aucubin content of seeds.] Sacar, G. R., & J. L. HARPER. Factors affecting the germination and early establishment of plantains (Plantago lanceolata, P. media and P. major). Pp. 236-245 in J. L. Harper, ed., The biology of weeds. Oxford. 1960 & . Controlled interference with natural populations of Plantago lanceo- lata, P. major and P. media. Weed Res. 1: 163-176. 1961. [Grasses affect size and eeu of Plantago populations, at least partially by acting on seedlings.] Biological flora of the British Isles[;] Plantago major L., P. media L. and P. lanceolata L. Jour. Ecol. 52: 189-221. 1964. [Extensive and detailed biological information; rosette clumping in P. /anceolata due to axillary shoot formation on underground rhizomes. SHapowsky, A. E. Embryological researches on Plantago major L. f. phyllostachya Wallr. (Russian summary.) Bull. Soc. Nat. Moscou II. 32: 234-259. 1924. [Pollen, SHEAR, G. M. Plantago arenaria. Claytonia 2: 14. 1935. [One individual of P. arenaria found near Blacksburg, Virginia. SHINNERS, L. H. Str ray notes on Texas Plantago Satie crams: Sida 3: 120-122. 1967. ———. Manual of the southeastern flora. xxii + 1554 pp. New York. 1933. [Nine spp. of ie from the area east of the Mississippi River in our range Smit, A. J., & J. W. WoLpeNporp. Nitrate production in the rhizosphere of Plantago species. Pl. Soil 61: 43-52. 1981. [P. major better adapted physiologically (levels of nitrate reductase) to nutrient-rich soils than is P. lanceolata. Situ, E. B. An atlas and annotated list of the vascular plants of Arkansas. iv + 952 pp. Published by author, University of Arkansas, Fayetteville, Arkansas. 1978. [Plantago, 251; nine spp. mapped, 252.] SnypDER, W. E. Mechanism of the photoperiodic response of Plantago lanceolata L., a long-day plant. Am. Jour. Bot. 35: 520-525. 1948 STEARNS, F. Effects of seed environment during maturation on seedling growth. Ecology 41: 221, 222. 1960. [P. aristata.] 1984] ROSATTI, PLANTAGINACEAE 561 STEBBINS, G. L., & A. Day. Cytogenetic a for long continued stability in the genus Plantago. Evolution 21: 409-428. 1967. [Remarkably little divergence has ace between Plantago ovata Cc Islands to India) and P. insularis (P. fastigiata) (southwestern United States and Baja California), in spite of over 20 million years of allopatry; cf., however, interpretation of BAssETT & BAuM; see also ASSETT & CROMPTON. ] STEINBAUER, G. P., & B. Gricssy. Dormancy and germination characteristics of the seeds of four species of Plantago. Proc. Assoc. Official Seed Analysts N. 158-164. 1957. [Primary dormancy nonexistent or of short duration in P. one and P. aristata; ae prevalent and persistent in P. major and P. Rugeli nee P., & A. D. J. Meeuse. Anthecological relations between nae ane- mo philous flowers and syrphid flies I. The nag role of syrphid flies as pollinators of Plantago. Tijdschr. Entomol. 119: 15-31. pls. 976. STEYERMARK, J. A. Flora of Missouri. 1xxxiii + ee pp. Ames, Iowa. 1962. [Plantago, 11 spp., 1380-1388.] STRAUSBAUGH, P. D. Branched spikes of Plantago Rugelii Decaisne. Castanea 15: 110. 1950 TeRAMURA, A. H. Experimental ecological genetics in Plantago IX. Differences in growth and vegetative reproduction in Plantago lanceolata L. (Plantaginaceae) from adja- cent habitats. Am. Jour. Bot. 70: 53-58. 1983. [Ecotypic variation, mostly with respect to irradiance, in ability to propagate vegetatively (clumping of rosettes due to axillary shoot production along underground rhizomes). ] , J. ANTonovics, & B. R. StrAIN. Experimental ecological genetics in Plantago IV. Effects of temperature on growth rates and reproduction in three populations of Plantago lanceolata L. (Plantaginaceae). Am. Jour. Bot. 68: 425-434. 1981. [Phys- iological adaptations to disturbance and temperate climates.] &B. TRAIN. Localized populational differences in the photosynthetic response to temperature and irradiance in Plantago lanceolata. Canad. Jour. Bot. 57: 2559- 2563. TESSENE, Me F. ‘Preliminary reports on the flora of Wisconsin no. 59. Plantaginaceae— plantain family. Trans. Wisconsin Acad. Sci. Arts Lett. 56: 281-313. 1968. [Exper- imental data used in character evaluation; a. information included for each species; consideration of P. major vs. P. Rugelii (capsules mistakenly said to be “dehiscent at middle’”’ in dean ton of the ae P. major “rarely found in hard- packed soils.” . Systematic and ecological studies on Plantago cordata. Michigan Bot. 8: 72- 104. 1969. THornE, R. F. The vascular plants of southwestern Georgia. Am. Mid]. Nat. 52: 257- 327. 1954, [Plantago, six spp., Trapp, G. A study of ie foliar endodermis in the Plantaginaceae. Trans. Roy. Soc. Edinb. 57: 523-546. 3. TRIVEDI, S., & R.S. ca Growth and reproductive strategies of two annual weeds as affected by soil nitrogen and density levels. New Phytol. 91: 489-500. 1982. [P. major.] The effects of soil texture and moisture on reproductive strategies of Spergula arvensis L. and Plantago major L. Weed Res. 22: 41-49. 1982. VANNEREAU, A., & J. C. MEstTRE. La région mer epylaite du sac embryonnaire du Diese ] latal «£¢0aaslt ) Revue Cytol. Biol. Vég. Bot. 3: 355-370. 1980. VerpcourRT, B. A new plantain from East Africa. Kew Bull. 23: 507-509. 1969. [Con- fusion between P. Cynops L., P. indica L., and P. Psyllium L Warwick, 8. I. The genecology ‘of lawn weeds VII. The response of different growth orms of Plantago major L. and Poa annua L. to simulated trampling. New Phytol. 85: 461-469. 1980. [Genetically determined prostrate growth forms of P. major 562 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 generally better able to withstand simulated trampling than corresponding erect growth forms . The genecology of lawn weeds III. Cultivation experiments with Achillea millefolium L., Bellis perennis L., Plantago lanceolata L., Plantago major L. and Prunella vulgaris L. collected from lawns and contrasting grassland habitats. Ibid. 83: 509-536. pl. J. 1979. [Habitat-correlated differences in plant size and erectness in P. lanceolata and P. major persisted (completely) in cultivation only in P. major, suggesting genetic control (cf., however, MARSDEN-JONES & TURRILL);, both genotypic differentiation and phenotypic plasticity important adaptive strategies in short-turf populations of P. /anceolata.] —- The genecology of lawn weeds. V. The adaptive significance of different growth habit in lawn and roadside populations of Plantago major L. Ibid. 85: 289- 300. 1980. [Genetically determined prostrate (from lawns) and erect (from roadsides) growth gen shown to be adapted to clipping and tall grass conditions, respectively.] Watts, W. A. e full-glacial vegetation of northwestern Georgia. Ecology 51: 17-33. 2 foldout a ee 1970. a reference to Wisconsin glaciation, Plantago pollen from pollen zones Q2-Q4 (probably late-glacial to early post- soe at Quicksand Pond and Q1 (probably full glacial) at Bob Black Pond, Bart WEATHERWAX, P. A variation in Plantago lanceolata. Proc. ere rer Sci. 1916: 365-367. 1917. WHITEHEAD, D. R., & E. S. BARGHOORN. Pollen analytical investigations of Pleistocene deposits from western North Carolina and South Carolina. Ecol. Monogr. 32: 347- 369. 1962. [Plantago pollen in Pleistocene sediments of uncertain age in the Carolina Pied : SS & D.J. READ. Vesicular-arbuscular mycorrhiza in natural vegetation s III. Nutrient transfer between plants with mycorrhizal interconnections. Nee Phytol, 90: 277-284. 1982. [Transfer occurred in cultivation experiments with P. lanceolata.] WopenHouse, R. P. Pollen grains. xv + 574 pp. New York and London. 1935. [Pollen of P. lanceolata and P. Rugelii illustrated, 451, Plantago oo P. lanceolata, P. major, and P. Rugelii) described and discussed, 455-457 Hayfever plants. xix + 245 pp. Waltham, Massachusetts. 1945. [Plantago, 125- 197 only P. lanceolata “sheds enough pollen and is itself abundant enough to cause hayfever.’ ] Wu, L., & J. ANTronovics. Experimental ecological genetics in Plantago I. Induction of roots and shoots on leaves for large scale vegetative ie ga and metal tolerance testing in P. ee New Phytol. 75: 277-282. erimental cic genetics in ane II. Lead tolerance in hs ne and Cynodon dactylon from a roadside. Ecology 57: 205-208. oa AURA, A. Genetical studies of plantains I[.] Genic analysis in the hexaploid and Hee plants. Bot. Mag. Tokyo 68: 35-39. 1955. [Branched spikes in P. japonica due to simple Mendelian recessiveness, but not expressed under very humid con- ditions. ] YOUNGKEN, H. W. A comparative study of the seeds and spikes of certain caulescent species of Plantago. Am. Jour. Pharm. 106: 157-165. 1934. [P. arenaria, P. Cynops, P. Psyllium.) BIOLOGICAL SURVEY New York STATE MUSEUM THE STATE EDUCATION DEPARTMENT ALBANY, New York 12230 1984] LAFRANKIE, STEM ABSCISSION 563 ANATOMY OF STEM ABSCISSION IN THE GENUS SMILACINA (LILIACEAE) JAMES V. LAFRANKIE, JR. ABSCISSION, the shedding of organs or parts, is a phenomenon found in many taxa of both higher and lower plants. The abscission zone associated with the leaves, fruits, and stems of dicotyledons of the Temperate Zone has received much attention, and comparative anatomical studies have been particularly extensive (Lee, 1911; Pfeiffer, 1928; Kozlowski, 1973; Addicott, 1982). An abscission zone typically includes two distinct anatomical regions: a discrete separation layer responsible for the actual detachment of the abscised organ; and a protective layer that seals the parent axis, preventing water loss and the entry of pathogens. While these two anatomical layers have been found in many dicotyledons, the occurrence of similar layers in monocotyledons is less well known. Sepa- ration layers have been found at the leaf insertion of some palms (La Floresta, 1904; Tomlinson, 1962) and in the fronds of the Lemnaceae (Witztum, 1974, Newton ef al., 1978). Protective tissues not necessarily associated with the separation of parts occur in many species of monocotyledons. Philipp (1923) classified the protective layers into three types: a metacutis, a simple periderm, and a storied periderm. Philipp, however, was not concerned with the condition or situation in which a plant developed protective layers, and it is unclear when—if ever—these layers are related to abscission. Stem abscission occurs in many monocotyledons of the Temperate Zone that grow sympodially. Such plants form a stem-abscission zone either at or below ground level as part of their normal seasonal cycle of shoot senescence and shoot regeneration. In this article, the developmental anatomy of abscission is described in Smilacina racemosa (L.) Desf. (Liliaceae), a good example of a sympodially regenerating herb with a perennial rhizome. For comparative purposes the abscission scars of four other species in the genus (Smilacina japonica A. Gray, S. stellata (L.) Desf., S. scilloidea Martens & Galeotti, and S. paniculata Martens & Galeotti) are examined, although in less detail. The annual shoot of Smilacina racemosa is determinate and consists of a horizontal rhizome and an “upright leafy axis. The aerial parts live only a single growing season; they juently replaced by a renewal shoot that develops from a lateral bud found at the base of the leafy axis. The renewal shoot grows horizontally for 2-3 cm before turning upward and forming an overwintering bud that contains the preformed leafy shoot for the following year. This se- quence of determinate growth and lateral regeneration is repeated annually. © President and Fellows of Harvard College, 1984. Journal of the Arnold Arboretum 65: 563-570. October, 1984. 564 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Although the aerial portion of each shoot is lost in autumn, the underground part lives for ten or more years; the perennial rhizome is thus a sympodium composed of the persistent basal portions of old shoots. A prominent scar marks the location of each of the abscised parent axes and therefore also the annual growth increment (FiGurE |). MATERIALS AND METHODS The abscission zone in Smilacina racemosa was studied in two ways: as it developed without disturbance in local populations at Harvard Forest, Peter- sham, Massachusetts; and as it developed in response to clipping of the aerial shoot. Previous experience showed that the abscission zone appears naturally during the late summer or fall prior to senescence of the aerial shoot, but developmental studies are difficult because the phenology of individual plants is quite variable. Abscission can be induced in midsummer by cutting the aerial stem just below the lowest leaf, approximately 25 cm above the ground. Such experimentally manipulated plants are advantageous because the abscission zone develops uniformly in them after the aerial stem has been cut. The aerial shoots of 25 plants within a natural population were cut in this manner on 27 July 1983, and the rhizomes of these plants were subsequently excavated, five at a time, at 0, 3, 7, 14, and 28 days after cutting. Untreated control plants were also collected on each occasion. Additional collections were made throughout the spring, summer, and fall of 1982 and 1983 to extend knowledge of the natural phenology of the protective process. Fresh material was examined microscopically in transverse and longitudinal hand sections. Additional material was fixed in FAA, dehydrated in TBA, embedded in paraffin, and sectioned at 8 um. Sections were either stained in safranin and alcian green or subjected to several microchemical tests. The presence of suberin was verified by Sudan IV and the IKI-sulfuric acid test (Johansen, 1940). Lignin was detected by both the HCl—-potassium perman- ganate test (Maule’s reagent) according to Johansen (1940) and the HCI-phlo- roglucinol reaction. In the latter method sections were first dehydrated in eth- Ficures 1-8. Smilacina racemosa, development of abscission zone. 1-3, 8, untreated control plants; 4-7, abscission induced; all sections stained using HCl-phloroglucinol method. 1, rhizome (August 30), x 0.9; abscission zone visible at base of aerial shoot (arrow), renewal shoot to left, scar of previous year’s aerial shoot to right. 2, thick longitudinal section through stem base (June 30), x 1.8, showing vascular bundles (vb). 3, thick longitudinal section through stem base (August 30), x 1.8, showing abscission zone (az). 4, longitudinal section through stem base (7 days after cutting), x pie showing first stage of abscission; some cell walls lignified. 5, t hrough stem base at level of abscission zone (7 days after cutting), x 75, showing aa See bundle before occlusion of tracheids (t). 6, pans verse section through stem base at level of abscission zone (14 days after itting), x 75, parenchyma cell walls heavily lignified and tracheids (t) occluded. i, Tong auclinel section through stem base (14 days 1984] LAFRANKIE, STEM ABSCISSION 565 et “ en <=] ) i after experimental cutting), x 80, showing newly divided cells proximal to ligno-su- berized layer. 8, longitudinal section through stem base (2 years after loss of aerial stem), x 80. 566 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 TasLe |. Voucher specimens. SPECIES VOUCHER Smilacina racemosa U.S.A., Massachusetts, Petersham, LaFrankie 138. S. stellata U.S.A., Massachusetts, Plum Is., LaFrankie 76. S. japonica Japan, Yonezawa (material in cultivation at Harvard For- est, Petersham, Massachusetts; no voucher collected). S. scilloidea Mexico, Oaxaca, Ixtlan, LaFrankie IV-21a. S. paniculata Costa Rica, Cerro del Muerte, LaFrankie IV-10c. anol series for complete extraction of soluble sugars and residues of decay, then vacuum infiltrated with 2 percent phloroglucinol in 95 percent ethanol and transferred directly to concentrated hydrochloric aci The living and preserved rhizomes of four other ee of Smilacina were sectioned and stained in a manner similar to that described above. Living plants of these species were not experimentally treated, and development of the abscission zone was not studied directly; however, by examining the suc- cessively older scars found within a single rhizome, I could draw some inference concerning the later modification of the abscission zone. Voucher specimens have been deposited in the Gray Herbarium, Harvard University (Gu) and are listed in TABLE 1. RESULTS SMILACINA RACEMOSA The parenchyma and vascular bundles at the base of the aerial stem are not distinguishable from the surrounding tissue during the spring and summer (FiGure 2). As individual stems begin to wither between July and October, a distinct abscission zone forms in the stem base at a scale-leaf internode 1-2 mm above the point of anatomical transition from the rhizome to the aerial stem (FiGures 1, 3). The rhizome has an endodermis, amphivasal vascular bundles, and isodiametric parenchyma cells; the aerial stem lacks an endo- dermis but has a peripheral fibrous sheath, collateral vascular bundles, and elongated parenchyma cells. The position and appearance of the abscission zone is the same whether it occurs naturally or is induce The abscission zone consists of an outer suberized and lignified layer and an underlying layer of dividing cells that later also become suberized and lignified. The aerial stem usually separates from the rhizome just above the outer ligno- suberized tissue, but there is no distinct separation layer, and often the dried stem remains weakly attached for a year or more. Three stages can be recognized in the development of the abscission zone. In treated plants the first is seen seven days after cutting. (Plants collected zero and three days after cutting show no changes associated with abscission.) In 1984] LAFRANKIE, STEM ABSCISSION 567 tothi natural (i. e., untreated ally coincides with the first yellowing of the leaves, but ihe: rate of eee varies considerably from plant to plant. In the first stage the primary walls of the cortical parenchyma cells become suberized and partially lignified (FicurE 4). The development of this ligno-suberized layer spreads inward to the center of the stem and outward until it includes the epidermis. The tracheids of most vascular bundles are water filled in fresh sections and are free of occlusions (FIGURE 5). The second stage appears in treated plants 14 days after cutting; in untreated plants it is found when the entire aerial axis has turned yellow or retains only patches of green, and when at least a few leaves have dried and withered. This stage is characterized by an intensified lignin reaction, the occlusion of some of the xylem elements, and divisions within cells proximal to the ligno-su- berized layer. The lumens of many tracheary elements are occluded with dark, amorphous contents (FIGURE 6), presumably gums or tannins, while other tracheary elements are air filled when examined in fresh sections. Cells im- mediately adjacent to the ligno-suberized layer repeatedly divide, with the new cell walls usually forming parallel to the plane of abscission. This results in subdivided cells (FiGuRE 7). There is no evidence of a phellogen (i.e., perma- nently meristematic cells). At this stage the walls of the dividing cells are neither suberized nor lignified. The third stage in stem abscission is first seen in the experimental plants 21 days after cutting but is found in untreated plants only when the main axis of the aerial shoot is dry and partially collapsed. In this stage the new walls of subdivided cells become suberized and lignified (FiGuRE 8). No new cell di- visions occur in adjacent cells. Xylem elements are completely occluded in all vascular bundles, and the occlusions extend 2 mm or more into the rhizome. In untreated plants the tissue distal to the ligno-suberized layer may decay, causing the withered aerial shoot to separate from the rhizome. Sometimes the weight of the terminal infructescence breaks the stem adjacent to the protective layer, but no mechanically weakened point or clear separation layer forms. As the aerial axis dries and withers, the ligno-suberized layer and the xylem oc- clusions become dark red-brown. The development of the protective layer is now complete; no further changes occur with age. OTHER SPECIES OF SMILACINA Two other species of Smilacina examined (S. stellata, S. japonica) have stem- abscission scars anatomically similar to those described for S. racemosa. There is a ligno-suberized layer of undivided and divided cells, beneath which occurs seemingly normal parenchyma. Smilacina paniculata, an herb from montane cloud forests in Costa Rica, differs from the other species studied in the greater width of its aerial stem, and in that the rhizome of the renewal shoot swells to several times its initial diameter soon after the aerial axis emerges (FIGURE 9). Consequently, the abscission scar in S. paniculata is much larger than that found in any of the other species. The rhizome of S. paniculata is also distinct in that the epidermis is lost during the initial expansion of the rhizome, and a simple periderm subsequently develops on all of the exposed rhizome surfaces (FiGuRE 10). 568 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 &. bab es of am Oe & pas , : Y FiGures 9-12. Smilacina paniculata, rhizome morphology and longitudinal sections. 9, rhizome viewed from above, x 0.2, showing scars of old shoots (arrow). 10, periderm of free rhizome surface, stained in safranin and alcian green, x 120. 11, stem base ca. 2 years after abscission of aerial shoot, stained in HCl-phloroglucinol, x 40; initial pro- tective layer (ip) above, vascular bundle (vb) ruptured by periderm (p). 12, lower region of abscission zone, x 120, showing dividing parenchyma cells adjacent to periderm, x 100 When the abscission zone of Smilacina paniculata first develops at the base of the aerial shoot, it is similar to that described for S. racemosa (1.e., a ligno- suberized layer of two or three cells). A few cell divisions occur in the paren- chyma on the rhizome side of the ligno-suberized layer, and the resulting cells are subsequently incorporated into the protective layer. The tracheids of the vascular bundles are occluded several millimeters into the rhizome. These features characterize the protective region for approximately one year after the loss of the aerial shoot. In older portions of the rhizome, a thick periderm develops beneath the ligno-suberized layer (FIGURE | 1), transecting the vascular bundles and augmenting the protective tissue of the abscission scar. The peri- derm of the abscission region joins with that of the free rhizome surface, but the two types of periderm can be distinguished by structural differences. The periderm of the free rhizome surface consists of radial files of six or 1984] LAFRANKIE, STEM ABSCISSION 569 seven cells that are uniform in size and shape, suggesting that they are deriv- atives of a phellogen, although this phellogen appears not to be permanent. When mature, all of the cells of the periderm are fully suberized and lignified and are quite distinct from the adjacent parenchyma. No cells occur in inter- mediate stages of development, which indicates that the periderm is not aug- mented through the continuous generation of new cells. The periderm of the abscission region is thicker than that of the rhizome surface, it is not clearly demarcated from the adjacent parenchyma, and there is no indication of a phellogen. The mature periderm cells are varied in size and shape (FicureE 12). Cell divisions seem to occur in parenchyma cells of progressively deeper levels, so that even when most of the periderm cells are suberized and lignified, parenchyma cells to the interior can be found dividing. Consequently, no eg line separates the periderm from the otherwise normal rhizome parenchym The other een species examined, Smilacina scilloidea, 1s similar to S. racemosa and S. stellata in general morphology, but it develops a periderm on its free rhizome surface and in the abscission-scar region; the periderm is anatomically similar to that found in S. paniculata. DISCUSSION The abscission zone of Smilacina consists only of a protective layer that seals the parent stem; no discrete separation layer is formed. Such a situation may be characteristic of geophytes with sympodial rhizomes, but this generality can be supported only if many more species are examined. The protective layer that seals the rhizome in Smilacina racemosa is notable for its early development, for the limited number of cells involved, and for its durability. In all but the last respect, the protective layer is similar to that found in the leaf-abscission zone of herbaceous dicotyledons. In plants such as Phaseolus and Gossypium, the initial protective layer forms when the cell walls of three or four layers of cells on the stem side of the separation layer become suberized and lignified. Tyloses occlude the vascular elements, and a few cell divisions occur adjacent to the ligno-suberized layer (Gawadi & Avery, 1950; Lee, 1911). Lee (1911) called these tissues the “‘initial protective layer”’ to distinguish them from a secondary protective layer that develops from a phellogen. It is quite remarkable that in S. racemosa such an initial protective region is able to seal the rhizome and remain intact for many years or even decades. Many woody dicotyledons augment the initial protective layer with a peri- derm that develops some distance to the stem side of the abscission zone. Lee (1911) described the various ways in which this can occur, but typically a phellogen develops below the initial protective layer and establishes continuity with the periderm of the parent stem. In Smilacina paniculata the initial protective region is likewise augmented by a periderm, but a phellogen does not develop. Rather, the periderm is derived through the periclinal (and oc- casionally anticlinal) divisions of subjacent cells, and in this respect it more closely resembles the wound tissue of monocotyledons described by Swamy 570 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 and Sivaramakrishna (1972) than it does either the periderm found in the rhizome surface or the periderm described for other monocotyledons (Philipp, 1923). Admittedly, periderm tissue is poorly known in monocotyledons, and a better appreciation of the relationship of wound tissue to periderm tissue awaits their more careful comparative analysis. The protective region described for Smilacina can best be considered as an anticipatory wound response, es- tablishing a water-tight seal prior to an actual break in the aerial axis. ACKNOWLEDGMENTS Monica Mattmuller and Frank Ewers suggested several improvements in microtechnique, William Ormrod assisted with the photography, and P. B. Tomlinson pointed out several important references and critically reviewed the manuscript. Specimens from Japan were generously provided by M. Ta- kahashi, while those from Costa Rica and Mexico were collected with the support of grants from the Atkins Garden Fund and the Anderson Fund of Harvard University. LITERATURE CITED AppicoTt, F. T. 1982. Abscission. University of California Press, Berkeley. Fioresta, P. ra. 1904. Sul meccanismo della caduta delle foglie nelle palme. Contr. Biol. Veg. 3: 255-273. Gawapl, A. G., & G. S. Avery. 1950. Leaf abscission and the so-called ‘‘abscission layer.” Amer. J. Bot. 37: 172-180. JOHANSEN, D. A. 1940. Plant microtechnique. McGraw-Hill Book Co., New York. Koz.owsk1, T. T., ed. 1973. Shedding of plant parts. Academic Press, New York. Lee, E. 1911. The morphology of leaf-fall. Ann. Bot. (London) 25: 51-105. Newton, R. J., D. R. SHELTON, S. DiISHAROON, & J. E. Durrey. 1978. Turion formation and germination in Spirodela polyrhiza. Amer. J. Bot. 65: 421-428. PreIFFER, H. 1928. Die pflanzlichen Trennungsgewebe. Jn: K. LinsBAueErR, e ae buch der Pflanzenanatomie. Band V. Lief. 22. Gebriider Borntraeger, Ber Puitipp, M. 1923. Uber die verkorkten Abschlussgewebe der Monokotylen. Poole Bot. (Stuttgart) 92: 1-28. Swamy, B. G. L., & D. SIVARAMAKRISHNA. 1972. Wound healing responses in mono- cotyledons. I. Responses in vivo. Phytomorphology 22: 305-324. TOMLINSON, P. B. 1962. The leaf base in palms: its morphology and mechanical biology. . Arnold Arbor. 43: 23-50. Witztum, A. 1974. Abscission and the axillary frond in Spirodela oligorhiza (Lem- naceae). Amer. J. Bot. 61: 805-808. HARVARD FOREST PETERSHAM, MASSACHUSETTS 01366 1984] INDEX INDEX Asse, Ernst C., and Rosert B. KAut_. In- florescence Architecture and Evolution in the Fagaceae, 375-401 Abelia dielsii, 60 Aberemoa, 216, 217, 231, 235, 240 Abies fargesil, 60 > ras o ae N Ne) 32 — — caudatum var. multiserratum, 60 — pensylvanicum, — tschonoskii, "60 ukurunduense, 60 hen. 216, 218, 235 Acioa, 216, 218, 235 Ackama, 168 Acouroa, 216, 230, 231, 235, 240 Acrophyllum, 150, 156, 164-166, 168 — australe, 151, 156, 158, 172, 173 Acsmithia, 149, 150, 155, 157, 164, 165, 9 es densilon: 151, 154, 157, ie 172-174 — undulata, — vitiense, Actaea asiatica, 60 — pachypoda, 60 frat melegueta, 19 Agalmyla, Agarista er A Taxonomic Revi- sion of the American Species of, 255- Agarista, 255-342 — sect. Agarista, 256-261, 264, 265, 269, 270, 273, 274, 282 — sect. Agauria, 256-258, 264, 265, 270, 273, 339 — sect. Euagarista, 282 — sect. Leucothoides, 282 — acicularis, 270 — albiflora, 259, 261, 264, 266, 267, 270, 271, 213; 2104 29, LOH 287,; 290 — ambi igua _ anastomosan s, 339 — angustissima, a 263, 266-272, 277, 334-338 — boliviensis, 261, 263, 266, 267, 270, 271, 277, 280, 328, 333-335 _ bracamorersis: 264, 266-268, 272, 279, 86 — bradei, 303 — breviflora, 325 — chapadensis, 263, 266, 270, 280, 302, 305-307 — chlorantha, 263, 266, 267, 270, 272, 274, 279, 282, 286, 313, 318-323, 325- =x igiliata. 939 — coccinea, 339 — coriifolia, 259, 261, 263, 267, 269, 271- 273, 278, 280, 281, 300, 303-305, 312, 316 — — var. bradei, 266, 271, 273, 294, 300, 302, 303 — — var. coriifolia, 266, 294, 300-303 — duartei, 261, 263, 266-270, 272, 277, 334, 338, 339 — duckei, 261, 264-267, 270, 271, 273, 279, 281, 284-287, 290 — ericoides, 261, 263, 266-273, 278, 286, 320, 324, 326, 327, 329 — criophylla, 339 — eucalyptoides, 259, 261, 263, 266, 267, 269-271, 273, 277, 280, 313, 328, 333- 335, 337 — glaberrima, 261, 266, 267, 269-271, 277, 334-338 — hispidula, 259, 261, 263, 266-268, 270, 272, 273, 278, 286, 313, 317, 320, 321, 323-327, 329 — ilicifolia, 339 S12 Agarista intermedia, 270 ei 9 - lycopodioides, 270 — mexicana, 265, 267-271, 280, 281, 285, 290, 293, 295, 299, 333 — — var. mexicana, 266, 290-293 — — var. pinetorum, 266, 270, 283, 290, 292, 293, 306, 307 — minensis, 266, 271, 272, 274, 278, 280, 308, 311-314, ae 317, 331 — multiflora, 334 — neriifolia, 300, 303 — miederleinii, 259, 261, 263, 268, 273, 278, 279, 281, 314, at. 329, 331, 332 — — var. deutifolia. 266, 273, 312, 328- 331 — — var. niederleinii, 266, 328-332 — nummularia, 263, 265, 266, 268, 270- 272, 274, 277, 282, 317- 321, 323 — oleifolia, 259, 261, 263, 267, 273, 280, 303, 307-309, 312-314, 316, 325, 332 — — var. glabra, 266, 270, 273, 308-312 — — var. hispidula, 270 — — var. oleifolia, 266, 270, 294, 308- 310, 312, 314 — organensis, 263, 266-268, 270-272, 278, 286, 320, 323-326, 329 — paraguayensis, 266, 269, 270, 273, 274, 279, 280, 306-308, 312 — populifolia, 257-259, 261, 263, 265- 272, 274, 275, 281, 285, 293, 295-300, 332 — pulchella, 265, 267, 270-273, 277, 278, 280, 307, 312, 314-317, 319, 323, 331 — var. cordifolia, 263, 266, 308, 312, ~315- 317, 331 — — var. eons 266, 303, 308, 311, 312, 314-317 — pulchra, 266-270, 277, 278, 280, 302- 306 — revoluta, 259, 261, 264, 265, 271, 273, 278, 279, 283, 285, 287-290 — — var. revoluta, 266, 288, 289, 302 — — var. velutina, 266, 270, 288-290, 302, 306, 307 — salicifolia, 256, 258, 264, 265, 270, 273, 274, 277, 339 — serrulata, 270, 282, 321, 323 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Agarista sleumeri, 265-271, 274, 281, 285, 292-295, 99 — spectabilis, 310 — sprengelii, 288 — stenophylla, 309 — subcordata, 261, 264, 266, 267-273, 278, 279, 284-286, 316 — subrotunda, 263, 266, 270, 277, 280, —307 — uleana, 266, 273, 281, 312, 330-333 — varnhageniana, 312 — virgata, 261, 263, 266-271, 277, 318- 321 Agauria, 255-257, 274, 339 — salicifolia, 274 ees 121 Alouea, 216 Aistopstalun, 158, ae fe 167 — multiflorum, 151, 4 —viticoides, 161 Allamanda, 223, 235, 241 Alliaria, 367, 373 — petiolata, 372 Allophylus, 108 — cobbe, 113 Alnus, Alpinia, 8, 19, 25 — galanga, | — officinarum, 16, 19 AL-SHEHBAZ, IHSAN A. The Tribes of Cru- ciferae (Brassicaceae) in the Southeast- 3 94 Alyssum, 347, 348, 350, 365, 371 Amaioua, 21 Amanoa, 21 ese ae ie 549 Amasonia, 228, 235 Ambelania, 216, 223, 233, 235 Amechania, 257, 274 — hispidula, 325 — subcanescens, 274, 321 American Species of Agarista (Ericaceae), A Taxonomic Revision of the, 255-342 Amyris elemifera, 114 Reo 108, 238 Anagallis, Anastatica a ochunties: 349 Anatomy of Stem Abscission in the Genus Smilacina (Liliaceae), 563-570 1984] Anatomy of the Palm Rhapis excelsa, X. Differentiation of Stem Conducting Tis- sue, 191-214 Anatomy, Vegetative, and The Taxonomic Status of Ilex collina and Nemopanthus (Aquifoliaceae), 243-250 -_ oe 333 — laurina, 297 _ longepetiolata, 334 7 — nitida, 289 | 5 Ss 3 5 =n bev] | i Ww x _ See 309 | o> j [o) co Pa =a E: Ow = & Ne) Ns) ~) pulchella, 2 15 — reticulata, 297 38 Anodopetalum, 158, 162-169 — biglandulosum, 151, Antirhea, 221, 235 Apalatoa, 216, 229, 235 Aquifoliaceae: Vegetative Anatomy and the Taxonomic Status of Ilex collina and Nemopanthus, 243-250 Aquifoliaceae, 233, 238, 243 Arabidopsis, 365, 373 INDEX 573 Arabis, sale hae 366, 371 canaden Azalia sre (Araliaceae), The Architec- ure of Devil’s Walking Stick, 403-418 Aralia, 403, 406, 412, 414-417 — spinosa, 403-418 Araliaceae: The Architecture of Devil’s Walking Stick, Aralia spinosa, 403-418 — and Evolution in the Faga- Inflorescence, aie Peainunaes of Devil’ Iki lia spinosa gies The, 403- ne Ardisia, 220, 232, 235 a, 217 Arrowroot Family, 39 Aruba, 217 Asarum caudigerum, 60 Ascium, 223, 235 Asclepiadaceae, 238 ee 220, 232, 235 Athyrium, 60 _ Goh aos 60 — fallaciosum, 60 — filix-femina, 60 Aubletia, 217, 231, 235 Aublet’s Generic Names by His Contem- age and by Present-Day Taxono- s, The Treatment of, 215-242 ree deltoidea, 350 Austrobuxus, 120 — carunculatus, 105, ig 117, 119 — germinans, 113 BAAS, Pieter. Vegetative Anatomy and the Taxonomic Status of Ilex collina and Nemopanthus (Aquifoliaceae), 243-250 Seis 217, 231, 235 Bagassa, 217 Baler 217, 239 Bana peters 367, 371 Barreria, 225, 233, 235 BARRINGER, itanthus, a New Kerry. Cub Genus of Gesneracenc from Brazil, 145- ue Bassovia, 217 eee “150. 165-167, 169 574 Bauera capitata, 150 Bellardia, 229, 234, 235 Bellonia, 217, 235 Benteca, 216, 235 Berberidaceae: A Monograph of Diphyl- leia, 57-94 Berberidaceae, 57, 67, 69, 70, 89 — subfam. Podophylloideae, 69 Berberis, 58, 68 Betula albo-sinensis var. septentrionalis, 60 — alleghaniensis, 60 , 60 Betilaceae: 375, 396 Bihai, 429, 431, 4 — subg. asia eam 431, 447 — subg. Taeniostrobus, 431, 447 — collinsiana, 482 — elegans, 494 — longa, 486 — marginata, 472 — platystachys, 480 Bockia, 222, Bocoa, 217 Bombacaceae, 109, 238 Bonnetia, 221, 232, 235 Bonnetiaceae, 231, 232, 238 BouFrrorD, Davin E., TsUN-SHEN YING, d Susumu TERABAYASHI. A Mono- graph of Diphylleia (Berberidaceae), 57- 4 Bougueria, 534, 536 5 Brassica, 345, 350, 366, 369 — campestris, 350, 368 igra, — oleracea, 350 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Brassicaceae: The Tribes of Cruciferae in the Southeastern United States, 343-373 Brassicaceae, 34 razil, Cubitanthus, a New Genus of Ges- neriaceae from, 145-147 ro wines i Broo G. L. WessTer, W. J. Havoen ae T. Giiuis, and D. E. STONE. Systematics and Palynology of Picro- dendron: Further Evidence for Rela- tionship with the Oldfieldioideae (Euphorbiaceae), 105-127 Brosimum, 219, 225, 235 Brucea, 120 Brunellia, 170 Brunelliaceae, 169, 170 Buchenavia, 224, 233, 235, 240 35 — , 225, 235 Burseraceae, 109, 234, 238 Burtt, B. L. The First Species of Stau- ranthera (Gesneriaceae) from New Guinea, with General Notes on the Ge- nus, 129-13 Cabomba, ean eae 238, 240 Cacalia adenostyloides, 60 — muhlenbergii, 60 — roborowskii, 60 Cacoucia, 217 Caesalpinia, 218, 235 Cakile, 345, 349, 363, 369 — edentula subsp. harperi, 368 Calathea, 40, 42, 43 — cna 43 ebrina, 43 Caldcuvia, 149, 150, 155-157, 165, 168, 22 elena ee 173 — brassii, 151, 3 — celebica, ae on — clemensiae, 151, 156, 173 — fulva, 151, — nymanii, 151, — paniculata, 151, 156, 157, 173 — paniculosa, 151, 155, 157, 173 — papuana, 151, 156, 173 — rosifolia, 151, 154, 155, 157, 158, 173 | 156, 157, 168, 172-174 — rufa, 15 Calepina, 363, 364, 369 — irregularis, California, Central, A Possible Magnolioid 1984] Floral Axis, from ey Upper Cretaceous of, 95-104 Calinea Calo, 218, 219, 228, 235 Sere ae 150, 151, 156-158, 173 Calycorectes, 218, 231, — bergii, Camax, 226, 235 Camelina, 365, 371 Cananga, 216, 217, 235 Caniram, 226 Canna, 7, 9, 17, 26, 29-39 — subg. Canna, 32 — — sect. Bialatae, 32, 34 — pelea 32 — compac — edulis, 2, 34, 35, 53 — flaccida, 30, 32- — — xX iridiflora, 33 — x generalis, 32-34 Cannaceae, Marantaceae, and Zingibera- ceae: The Zingiberales in the Southeast- ern United States, 5-55 Cannaceae, 5-7, 9, 29, 31, 34, 39, 41-43, Capparaceae, 108, 238, 348, 349 — subfam. Cleomoideae, 346 Capparis, 230, 235 Caprifoliaceae, 534 Capsella, 364, 369 — bursa-pastoris, 368 Caraipa, 217, 231, 235, 239 — parvifolia, 231 Carapa, 217 Carapichea, 218, 219, 228, 235 Cardamine, 344, 345, 347, 349, 366, 371 — chenopodiifolia, 349 — clematitis, 60 Carinavalva, 346 INDEX Loishoglia bettencourtii, S75 ae 223, 233, 235 s, 2 Cassipourea, 218, 231, 235 , 398 Castanopsis, 375- 377, 383, 385, 386, 391- 396, 398 — acuminatissima, 385, 398 — ae — armata, 398 — ar 386, 398 — chevalieri — chrysophylla, 376, 386, 391, 392, 398 lifolia, 398 — lo ongipetiolata, 399 — megacarpa, 399 — motleyana, 399 — nephelioides, 399 — philippinensis, 399 Catalpa punctata, 113 Catinga, 218, 231, mee 239, 240 — moschata, 231, 24 Caulophyllum, 57 — robustum, 60, 65 Celaenodendron, 122 576 meee 232, 238 Celosia, 549 Gales oxymitrum, 18 Central American Holiconia (Heliconi- aceae) with Pendent Inflorescences, Sys- tematics of, 429-532 Centunculus, 549 35 Ceratopetalum, 158, 159, 163, 165-168 — succirubrum, 151, 159, 160, 172 Cercidiphyllum, 57 Chaetocarpus, 225, 233, 235 235 Chionanthus, 222, 232, 235 Chlora, 223, 235 Chomelia, 221, 235 Chorispora, 363, 373 hromosomes and Relationships of Meta- sequoia and Sequoia (Taxodiaceae), The: an Update, 251-254 Chrysobalanaceae, 231, 238 Chrysolepis, 376 Chrysosplenium microspermum, 60 Cimicifuga foetida, 60 6 Circaea alpina subsp. alpina, 60 — — subsp. imaicola Clayton, Temple, 1-4 Clayton, Temple, Chemist and Amateur Botanist, 1914-1978, 1 Clibadium, 217, 2 Clintonia udensis, 60 ulata, 60 Coccocypselum, 229, 233, 235 Codia, 159, 160, 163-169 , 159, vt 172, 174 Conohoria, 218, 232.933, 395: 240 Conopea, 218 5 JOURNAL OF THE ARNOLD ARBORETUM [OL 05 Conoria, 218, 235 Cornaceae: Some Observations on the Re- productive Biology of Three Species of Cornus, 419-427 Cornaceae, 419 Cornus (Cornaceae), Some Observations on the Reproductive Biology of Three Species of, 419-427 Cornus, 419-427 — subg. Thelycrania, 57 — florida, 419-426 — mas, 419-426 — sericea, 419, 421-426 ase 344, 364, 369, 542 — didymus, 368 Oe 3 7, 16, 433, 441 Costus, 7, - Coublandia, 218 Couepia, 26, 218, 231, 235 Couma, 218 Coumarouna, 219, 228, 235 Coupoul, 219, 232, 235, 240 Coutoubea, 219, 239 Craibiodendron, 255, 256, 261, 272 9 9 Cretaceous. Upper, of Central California, A Possible Magnolioid Floral Axis, Loishoplia bettencourtii, from the, 95- 104 Cruciaceae, 345 Cruciferae (Brassicaceae) in the Southeast- ern United States, The Tribes of, 343- 373 aber 343-373 tribe Alysseae, 346, 347, 363, 369-371 ae Anchonieae — tribe Arabideae, 347, 363, 370-372 — tribe Brachycarpeae, 369 — tribe Brassiceae, 343, 346, 347, 362, -370 — tribe Cakilineae, 369 — tribe Calepineae, 369 1984] Cruciferae tribe Camelineae, 371 — tribe Heliophileae, 346 — tribe Hesperideae, 346, 347, 363, 371- 373 — tribe Iberideae, 369 — tribe Isatideae, 369 — tribe Lepidieae, 343, 346, 347, 362, 368- 370 — tribe Lunarieae, 346, 371 — tribe Matthioleae, 346, 371 69 — tribe Romanschulzieae, 346, 367 — tribe Schizopetaleae, 346, 371 — tribe Senebiereae, 369 — tribe Sisymbrieae, 346, 347, 363, 372, 373 — tribe Stanleyeae, 346, 367 — tribe Stenopetaleae, 346 — tribe Streptantheae, 346, 367 — tribe Subularieae, 369 — tribe Thelypodieae, 343, 346, 347, 363, 367, 369, 370 — tribe Tinlaspideac: 369 — tribe Velleae, 369 Cubitanthus, A New Genus of Gesneri- ae from Brazil, 145-147 eae 145-147 — alatus Cucullaria, 230, 235 Cunninghamia, 221, 232, 235 Cunonia, ae 154, 155, 158, 163, 165, 168 — atrorubens, 151 — balansae, 152, 154, 174 — capensis, 152, 154 — lenormandii, 152 — macrophylla, 152, 156, 157, 172 — pulchella, 152, 154, 158, 172 INDEX eve Cunonia purpurea, 152, 156, 157, 172 — schinziana, 152, 157, 173 Cunoniaceae, Fruits and Seeds of the, 149- 190 Cunoniaceae, 149-190 —t 8 — tribe Spiraeanthemeae, 167 Cupressus, 29 Curcuma angustifolia, 19 — domestica, | — longa, 19 Cyclas, 216, 229, 235 Cyperaceae, 17, 233, 238 Cyrtandra, 130 30 Ga 225, 235 Dahuronia, 222, 235 Dalbergia, 216, 231, 235 Degeneria, 102 eee 102, 103 Deguelia, 219 Derris, 219, 2 Descurainia, 365, 373 — pinnata, Devil’s Walking Stick, Aralia spinosa (Ara- liaceae), The Architecture of, 403-418 Dialium, 217, 23 Dichapetalaceae, 238 DICKISON, WILLIAM C. Fruits and Seeds of he Cunoniaceae, 149-190 Dilleniaceae, 238 Dimerocostus, 16 imorpha, 219, 224, 235 imorpho a, 344 Diospyros, 224, 226, 235 oe Ape eae A Monograph of, 57-9 a ne 57-94 mosa, 57-78, 93, 94 — — subsp. grayi, 82 — — subsp. sinensis, 78 — grayi, 57-71, 78, g2- 89, 93, 94 — — var. incisa, 82, 89 var. rotundata, 82, 89 — — var. typica, 82 578 Diphylleia sinensis, 57-62, 64, 65, 67-71, -81 Diphylleiaceae, 69 Diplotaxis, 349, 366, 369 Draba, 345, 347, 350, 363, 365, 371 Dracontium, 226, 235 — sinofibrillosa, 60 Dugortia, 224, 2 Duguetia, 216, — 235 Dupinia, 228, Duroia, 219, a 235, 240 Dysosma, 64, 69, 70 Ebenaceae, 233, 238 Ensete, 432- 434, 448 Eperua, 219, 224, 235 Ephielis, 221, 235 Epithema peer 130 Ericaceae: A Taxonomic Revision of the merican species ce Agarista, 255-342 Spi 255, 258 e Andromedeae: 255, 269, 274 ar ie subsp. sativa, 370 Erucastrum, 36 Erysimum, 347, 348, 365, 373 0 trys, 257 Eucryphiaceae, 169 Euphorbiaceae: Systematics and Palynol- ogy of Picrodendron— Further Evidence for Relationship with the Oldfieldioi- deae, 105-127 Euphorbiaceae, 238 109, 114, 119-122, 233, JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Euphorbiaceae subfam. Oldfieldioideae, 105, 114, 116-122 — — tribe Acalypheae, 122 — — tribe Hyaenanchea — subtribe Dissiiarinae: 122 — — — subtribe Paivaeusinae, 122 — — tribe Picrodendreae, 122 Evea, 218, 219, 228, 235 Evolution in the Fagaceae, Inflorescence Architecture and, 375-401 Fagaceae, Inflorescence Architecture and Evolution in the, 375-401 Fagaceae, 121, 375-401 — subfam. Castaneoideae, 376, 377, 395 — subfam. Fagoideae, — subfam. Quercoideae, 376 — subfam. Trigonobalanoideae, 376 Fagus, 57, 376, 377 — crenata, 60 — grandifolia, 60 Faramea, 219, 235, 239 Ferolia, 219, 224, 235 First Species of Stauranthera (Gesneri- aceae) from w Guinea, The, with General Notes on the Genus, 129-133 Hae eT 234, 238 Foleyola, 349 Fothergilla, 220, 232, 234, 235, 241 Fraxinus, 2 Freyeria, 222, 235 Fruits and Seeds of the Cunoniaceae, 149- 190 Galarips, 223, 235 , 155, 157, 158, 163- — benthamiana, 152, 157, 172 — biagiana, 152, 156, 157, 172 — hippocastaneifolia, 152, 154 — pruinosa, 152, 154 — stipularis, 152, 157, 158, 172 Genipa, 223, 229, 235 1984] Gentiana, 223, 235 Gentianaceae, 232, 238 Geococcus pusillus, 349 Gerbera, Gerberia, 225, 236 Gesneriaceae from Brazil, Cubitanthus, a New Genus of, 145-147 Gesneriaceae: The First Species of Staur- cone from New Guinea, with General n the Genus, 129-133 Gesneriacee, 145, 146, 535 — tribe Beslerieae, 146 Gai 228, 236 Gillbeea, 159, 162-167 — papuana, 152, 172 GILLis, W. T., D. E. STONE, C. R. BROOME, L. Wester, and W. J. HAYDEN. Sys- tematics and Palynology of Picroden- dron: Further Evidence for Relationship with the Oldfieldioideae (Euphorbiace- ae), 105-127 Ginannia, 224, 236 Ginger Family, 15 Glaucidium palmatum, 59, 60 Globulariaceae, 535 Glossanthus grandiflorus, 131 Glossoma, 230, 23 Glossopetalum, 220, 236 Griselinia, 223, 236 Guapira, 220 Guatteria, 216, 217, 231, 236 UNATILLEKE, C. V. S., and I. A. U.N. me One. on the Reproductive Biology of Three ee of Cornus areal 419-42 GUNATILLEKE, I. A. U. N., and is V_S. GUNATILLEKE. Some Observations on the Reproductive Biology of Three a of Cornus (Cornaceae), ae Gustavia, 225, 233, 236, 24 Guttiferae, 238 Haemodoraceae, 238 Halopegia, Haloragaceae, 231 Hamamelidaceae, 232 INDEX 579 Hambergera, 217, 236 Hamelia, 227, 236 HAYDEN, W. J., W. T. GILiis, D. E. STONE, . R. BRoomME, and G. L. WEBSTER. Sys- tematics and Palynology of Picroden- dron: Further Evidence for Relationship with the Oldfieldioideae (Euphorbiace- ae), 105-127 Hedychium, 8, 17, 19, 21-29, 35 23, — coronarium, 7, 16, 21-26 — elwesu, 23 — flavescens, 23 — flavum, 2 _ gardnerianum, 26 Heliconia (Heliconiaceae) = Pendent Infloresce nces, Systematics of Central 32 mys, 43 — se ect. Taeniostrobus, 430, 431 — alleni, 478 — bihai, 429, 430, 461 — catheta, 480, 482 — colgantea, 436, 442, 444-446, 450, 454, 459, 462, 469-472, 524 — collinsiana, 438, 439, 442, 445, 446, 450, 454, 456, 462, 482, 483 — — var. collinsiana, 435, 450, 483-486, 52 — — var. velutina, 450, 480, 483-486 — curtispatha, 432, 435, 441, 444-447, 450, 456, 458, 459, 463, 486-490, 492, 516, 524 —-—*x ecm pogonantha, 515 — var. holerythra, 515, 516, 518 580 Heliconia danielsiana, 441, 444-446, 450, 456, 457, 463, 507-510, 512, 514, 524 — dresslerana, 512 — elegans, 494 _ imbricata, 430, 439, 447, 450, 451 — indica, 44 — irrasa a, 47 78 _ latispatha, 447, 450, 451 — longa, 486, 4 — maculata, 435, 436, 439, 442, 450, 454, 456, 462, 464-469, 524 — magnifica, 435, 436, 441, 450, 456, 457, 463, 505, 509-512, 514, 5 — marginata, 442, 450, 454, 456, 458, 462, 472-474 — mariae, 430, 432, 439, 441, 445-447, 450, 456-458, 463, 494-497, 516, 524 — x nagar pogonantha, 515, 518 = r. holerythra, 515, 516, 518 — mathiasiae, 447, 450, 451, 458 — metallica, 436 — x aiooreani: 483 — necrobracteata, 442, 450, 454, 456, 462- 466, 468, 524 _ nigripraefixa, S15 — nutans, 442, 443, 445, 446, 450, 454, 462, 475- 480, 524 — — x Heliconia ae 515, 518 — = X = — var ra, — platystachys, 434, 442, 450, 454, 456- 458, 462, 480-483, 489, 524 — pogonantha, 441, 443-446, 448, 450, 456-458, 463, 492, 497, 498, 501, 502 — — var. holerythra, 450, 498, 500, 501, 516 — — var. pogonantha, 450, 498-500 — — var. pubescens, 450, 498, 501 — — var. veraguasensis, 450, 498, 499, 501, 502, 524 — punicea, 432, 494 — ramonensis, 435, 441, 450, 456, 457, 463, 498, 502, 503, 505, 506, 512, 514 — — var. glabra, 450, 503-506 — — var. lanuginosa, 450, 503-507, 524 — — var. ramonensis, 450, 503-505, 507 — — var. xanthotricha, 450, 503-507, 510 — reticulata, 435 — rostrata, 482, 486, 487 — schiedeana, 483 — secunda, 436, 442, 447, 451, 454, 462, 473-475, 479, 524 — — x Heliconia clinophila, 515 — — var. secunda, 451, 474-476 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Heliconia secunda var. secunda x Heli- conia ers 51 _ r. viridiflora, 435, 439, 451, 474- 416 — sessilis, 436, 451, 456-458, 463, 492- — stilesii, 436, 439, 441, 443-446, 451, 456, 459, 463, 488-492, 524 — — x Heliconia danielsiana, 515, 518 — talamancana, 436, 442, 451, 454, 462, 466, 468, 469, 524 — tortuosa, 447, 451, 475, 478 — — x Heliconia nutans, 515, 518 — trichocarpa, 435, 438, 439, 442-446, 451, 454, 459, 462, 470-472, 524 — — var. caducispatha, 470, 472 — — var. trichocarpa, 472 — vellerigera, 505, 507, 510 , 45 — xanthovillosa, 436, 441, 451, 456, 457, 463, 509, 512-514, 524 Heliconiaceae: Systematics of Central American Heliconia with Pendent Inflo- rescences, 429-532 Heliconiaceae, 5, 432-434, 448 Heliconiopsis, 444, 461 — amboinensis, 461 Heliotropium ternatum, 113 Hesperis, 347, 365, 371, 373 — matronalis, 350, 372 Hevea, 220 Heymassoli, 220 Himantandraceae, 102, 103 Hirtella, 227, 236 Holacantha rtea, 170 Humiria, on 236, 240 Humiriaceae, 238, 240 Hutera, 366, 369 Hydrolea, 227, 236 Hydrophyllaceae, 238 Hypselodelphys, 50 Iberis, 348, 350, 363, 364 Icacinaceae, 234, 238 1984] Icacorea, 220, 232, 236 Icica, 220, 239 Idesia, 226, 234, 236 Ignatia, 226, 236 Ilex collina and Nemopanthus (Aquifoli- aceae), Vegetative Anatom the Taxonomic Status of, 243-250 Ilex, 221, 236, 243, 245, 249 — subg. Prinus, 243, 248, 249 — collina, 243-246, 248, 249 = verticillata , 249 Inflorescence ecniectune and Evolution , 375-401 Ivira, 22 Ixora, 224, 236 Japarandiba, 233, 236, 241 3 JouHNSON, L. C., S. E. SCHLARBAUM, and T. TsucHIYA. The Chromosomes and Re- lationships of Metasequoia and Sequoia (Taxodiaceae): an Update, 251-254 Joncquetia, 228, 236 Jupp, WALTER S. A Taxonomic Revision of the American Species of Agarista (Er- icaceae), 255-342 Juglandaceae, 106, 109, 120, 121, 396 Juglans baccata, 106, 108-110 Juncaceae, 17 Juniperus virginiana, 544 Kaempferia, 19 Kalawael, 234, 236 KauL, Ropert B., and Ernst C. Asse. In- florescence Aechitectiire and Evolution 01 Kress, W. Jon. Systematics of Central merica ni with Pendent Inflorescences, 429-532 Kruegeria, 230, 236 INDEX ee oe) 225; D222 LAFRANKIE, JAMES V., JR. Anatomy of Stem Abscission in the Genus Smilacina (Lil- iaceae), 563-570 Laguncularia racemosa, 113 Lagunezia, 226, Lamanonia, 7 a 157, 165, 167 Lecythidaceae, 238 Legnotis, 218, 231, 236 Leguminosae, 232, 233, 238, 240 6 32, 236 Lepidium, 344-347, 349, 364, 369 — campestre, 368 — sativum, 35 — virginicum, 368 Lesquerella, 344, 347, 365, 371 é 8 — ambigua var. ambigua, 309 — — var. glabr. — e — — var. tomentella, 309 — andina, 2 — angustissim a, 337 — bahiensis var. arrabidae, 289 582 eens bahiensis var. bahiensis, 288 r. bla 8 nchetil — var. canines 288 bracamorensis, 285 chlorantha, 321 — var. subcanescens, 321 var. laxiflor ra, 30 — var. martii, 301 — var. sellow1i, 300 oleifolia, as — — var. glab a, 310 — var. hispidala, 270, 309 — var. longifolia, 310 organensis, 323 paraguayensis, 307 a a, 3 pistrix, 305 JOURNAL OF THE ARNOLD ARBORETUM Leucothoé pohli, 301 — — var. cordifolia, 316 — pulchra, 304 — — var. parvifolia, 304 — revoluta, —_ Hodats 309 a — varnhageniana, 312 —_ vencalensis, 286, 287 Lew cae 220, 222, 231, 236 Licaria, 220 Lignonia, 224, 236 Ligularia stenocephala, 60 [voL. 65 Liliaceae: Anatomy of Stem Abscission in 0 the Genus Smilacina, 563-57 Liliaceae, 563 Limborchia, 219, 236 Lindera benzoin, 60, 416 — umbellata, Linociera, 222, 232, 236 — bumelioides, 113 Liquidambar, 291, 292, 295, 299 Liriodendron, $7, 100, 101, 103 — aggregata, 378, 380, 399 — amygdalifolia, 380, 399 — cantleyan — a, 399 — seen 381, 383, 399 2305 230 Lithocarpus, 376-378, 380-384, 391-396, 399 1984] Lithocarpus cornea, 399 — curtisii, 382, 384, 399 — pee 378, 399 — dealba — havilandii, 380, 399 — hendersoniana, 399 — hystrix, 399 - Eawakaniii: 384, 399 — kodaihoensis, 382, 399 — konishii, 399 — lampadaria, 383, 399 — leptogyne, 399 — lucida, _ macphaili, 399 — polystachya, 400 r — reinwardtii, 400 — wallichiand. 400 — woodil, 4 — wrayi, 400 Lithodraba oo 349 mye oe Pp na, an — Beis. 534 399 eine 376, 381, 391, 392, 399 INDEX 583 Littorella uniflora, 534, 535 ees. 234, 236 Loganiaceae, 234, 238, 535 Loghania, 227 Loishoglia bettencourtii, from the Upper Cretaceous of Central California, A Pos- sible Magnolioid Floral Axis, 95-104 0 — subg. Eubotrys, 257 6 Lythraceae, 238 Mabea, 220, 239 Maconcous 221, 233, 236 Macrolobium, 223, 230, 236 Magnolia, 57, 100-103 hoglia bettencourtti, from the eeey e Malcolmi, 363, 365 — afric Mee 301, 239 584 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Managa, 221 Menispermaceae, 238 Manettia, 223, 236 Metasequoia and Sequoia (Taxodiaceae), Mapania, 221 The Chromosomes and Relationships of: Mapouria, 221, 224, 227, 236 an Update Mappia, 227, 234, 236 Metasequoia, 251-253 Maprounea, 221 — glyptostroboides, , 253 Maquira, 221 Mexico, Notes on Salvia (Labiatae) in, with Maranta, 9, 40, 43, 51-55 Three New Species, 135-143 — subg. Automaranta, 52 Michelia, 100, 102, 103 — subg. Calatheastrum, 52 Miconia, 220, 232, 236 — subg. Friedrichsthalia, 52 Miegia, 226, 236 — subg. Koernickea, 5 Minquartia, 222 — subg. Maranta, 52 Mischodon, 120 — sect. Eumaranta, 52 — zeylanicus, 106, 119 — sect. Saranthe, 52 Mnasium, 226, — sect. Stromanthe, 52 Moelleria, 220, 236 — sect. Xerolepis, 52 Moniera, 231, 236 — arundinacea, 34, 40, 43, 51-54 Sauer eae 238 — bicolor, 53 Monocostus, 16 — divaricata, 52 eeiciars of Diphylleia (Berberidaceae), — aaedvichetaeliands 52 A, 57-94 — leuconeura, 53 Monophyllanthe, 52, 53 — — var. erythroneura, 53 Montira, 222 — ruiziana, Moauilea, 222 Marantaceae, Zingiberaceae, and Canna- Moraceae, 238 ceae: The Zingiberales in the Southeast- Morisia monanthos, 349 ern United States, 5-55 Moronobea, Marantaceae, 5-7, 9, 34, 39-45, 49, 50, Mourera, 222 433 Mouriri, 222 — tribe Maranteae, 40 Mouroucoa, 222 e Phrynieae, 40 Moutabea, 222 Marantochloa, 53 Moutouchi, 223 Marcgraviaceae, 238 Muellera, 218, 236 Marica, 218, 236 Musa, 8, pa 432-434, 448 Maripa, 221, 222, 236 — bihai Mariscus, 226, 233, 236, 240 ah 5, 8, 17, 43, 432-434, 448 Matayba, 221 — subfam. pe 433 Matelea, 221, 239 — subfam ideae, 433 Matourea, 221 — subfam. — 433 Matthiola, 363, 365 — — tribe Strelitzieae, 433 — incana, ai — tribe Heliconieae, 433 — longipetala, 363 — tribe Museae, 433 Matcha, 224, 236 Mussaenda, 227, 236 Mayaca, 222 Mustard Family, 344 aceon 238, 240 Myristica, 230, 236 Mayepea, 222, 232, 236 Myristicaceae, 238 Mayna, 222 Myrmecia, 227, 236 ebony 222 Myrodendrum, 220, 236 Megacarpaea megalocarpa, 349 Myrodia, 225, 233, 236 — polyandra, 34 Meisteria, 223, 236 Melastoma, 220, 221, 229, 236 Myrsine, 226, 236 elastomataceae, 232, 238 Myrtaceae, 238 Meliaceae, 238 Myrtus, 222, 236 1984] INDEX 585 Nacibea, 223, 239 Pachira, 223 Napimoga, 2 Soa 216, 223, 236, 240 Nasturtium, 367, 371 Pacourina, 3 Nemopanthus Ps erapeney Nee Anatomy and th c Status of Ilex collina and, te 50. Nemopanthus, 243, 248, 249 — collinus, 243 — mucronatus, 243, 245-249 New Guinea, The First Species of Staur- anthera (Gesneriaceae) from, with Gen- eral Notes on the Genus, 129-133 Nicandra, 225, 233, 236 2 Notes on Salvia (Labiatae) in Mexico, with Three New Species, 135-143 Nyctaginaceae, 238 Nyssa, 299 Ochnaceae, 238 Oc africana, 106, 118 Oldfieldioideae (Euphorbiaceae), System- atics and Palynology of Picrodendron: Further Evidence for Relationship with the, 105-127 Oleaceae, 238 Ouratea, 223, 232, 236, 240 Ourouparia, 223 Outea, 223, 230, 236 Oxalis acetosella, 60 Ozophyllum, 229, 236 Pagamea, 223 PAGE, VIRGINIA M. A Possible Magnolioid oral Axis, Loishoglia bettencourtu, from the Upper Cretaceous of Central California, 95-104 Palicourea, 224, 227, 236 Palm Rhapis excelsa, Anatomy of the, X Differentiation of Stem Conducting Tis- sue, 191-214 Paloue, 224 Palovea, 224, 236 Palynology of Picrodendron, Systematics and: Further Evidence for Relationship with Oldfieldioideae (Euphorbiace- ae), 105-127 Pamea, an 233, 236, 240 Panax pseudo-ginseng var. bipinnatifidus, Pancheria, 150, 155, 156, 158, 165, 168 — confusa, 152, 154, 155, 158, 172 — a 152, 154, 174 a, 60 Parivoa, 319, 224, 236, 239 Parolinia, 350 Passifloraceae, 238, 240 Passoura, 224, 226, 236 eats 224, 929, 236 Patim Pauline, 219, 236 Pavetta, , 236 Pecheya, 219, 236 Pekea, 224, 227, 236, 239 Peracarpa carnosa var. circaeoides, 60 Petaloma, 222, 236 Petrocarya, 224, 236 586 JOURNAL OF THE ARNOLD ARBORETUM Phaeomeria, 19 Phaseolus, 569 Saleen 432-434 Phrymaceae, 534 Plans 218, 222, 236 Physar 47 Picea is 60 bens, 60 36 Picrodendracca, 109, 1 2 ] Picroden 1 Palynology of: sis Evidence for Relationship with the Oldfieldioideae (Euphorbiace- -127 ae), 1 Picrodendron, 105-127 — arboreum, 108, 113 — baccatum, 106-116 — — var. bahamense, 108, 110 — juglans, 108, 110 — macrocarpum, are 110 — medium, 108, 110 Pieris, 255, 256, 272 Pinus, 291, 292, 295, 299 | io) ° ret 5 et 22 — longepedunculata, 106, 118 Piratinera Pirigara, 295. 233, 236, 239-241 — tetrapetala, 233 Plagiogyria matsumureana, 60 Plantaginaceae in the Southeastern United States, The, 533-562 Plantaginaceae, 533-562 Plantago, 534-536, 540, 542-562 — subg. Coronopus sect. Coronopus, 548, 550 t. Maritima, 551 — sb Plane 0, 543 [VOL. 65 Plantago subg. Plantago sect. Virginica, 545 — sect. Gnaphaloides, 546, 548, 550 — ser. Gnaphaloides, 546, 547 — sect. Lancifolia, ae 548, 550 sect. Montana, sect. Psyllium, 7 548 5 — sect. Leucopsyllium, 545-548 — sect. Novorbis, 545 — sect. Oreophytum, 545 — altissima, fey — arenaria — aristata, 7. 546, 547, 550 — cordata, 544, 545 — coronopus, 5 x Plantago eriopoda, 545, 550 s, 548 ~ lanceolata, ay 540, 542, 546-553 acro — major, 535, 34 -544, 547-550, 552, 553 — psyllium, 547, 549, 553 — purshii, — pusilla, 543, 544, 549 — rhodosperma, 545, 546, 550 _ rugelii, 541, 543, 544, 548-550 , 34 oe a 547, 550 Plantai Pl ack ‘Family , 533 Platylophys, 159, 162, — 165, 166, 169 714 — trifoliatus, 152, 172, Plotia, 226, 237 Plum baginaceae, 534-536 Podophyllaceae, 59, 69, 70 — subfa m. Glaucidioideae, 59 t. Lamprosantha, 550 _ ies Micropsyllium, 543, 546 — sect. Palaeopsyllium, 544, 545 — sect. Plantago, 543, 548, 550 — subfam. Podophylloideae, 59 aie ane 57, 59, 64, 69, 70 — peltatum, 59, a el gunaeeel 238 1984] Polygalaceae, 238 Polygonatum pubescens, 60 — roseum Polystichum acrostichoides, 60 — retrorso-paleaceum, Posoqueria, 22 Possible Magnolioid Floral Axis, Lois- hoglia bettencourti, from the Upper Cretaceous of Central California, A, 95- 104 Possira, 225 teria, 225 Sennen 534, 535, 549 Pringlea antiscorbutica, 349 20, 237 Prunus gracilifolia, 60 — grayana, Pseudanthus, 120 Pseudo- Brasilium, 228, 237 Pseudocarpidium wrightii, 113 Sg uauen ae spinulosa, 60 eudoweinmannia, 159, 160, 164-168 reese 152, 160, 172, 174 Psychotria, 218, 221, 223, 226, 228, 237 Psylliaceae, 534 Psyllium, 542 Pterocarpus, 223, 232, 237 Pullea, 159, 160, 163-169 — glabra, 152, 159, 160, 174 — var. glabra, 159, 160 — stutzeri, 152, 160, 174 Qualea, 225, 239 Quassia, 121, 227, 237 Quebitea, 226 Quercus, 291, 292, 295, 299, 376. 377, 381- 383, 386-395, 400 — acuta, 388, 400 _ borealis, 390, 400 INDEX 587 Quercus brandisiana, 389, 400 — cambodiensis, 389, 400 — championii, 388, 389, 400 — chapensis, 389 | le) a a 2 oO ue} a yn b io) oO — kinabaluensis, 389, 400 — kingiana, 389, 392, 400 — merrillii, 400 — miespilifolioides: 389, 400 — moril, 388, _ eee 388, 400 pachylom 0 paucident 400 - ees 388, 400 — salicina, 400 — serrata, 389, 400 stenophylloides, 388, 400 subsericea, 4 — uraiana, 386, 400 — valdinervosa, 389, 400 — virginiana, 400 Quiina, 226 Quiinaceae, 238, 240 Racaria, 226 , T. P. Notes on Salvia (La- biatae) in Mexico, with Three New Species, 135-143 Ramspekia, 225, 237 Ranunculaceae, 70 Rapistrum, 363, 369 Raputia, 226 Ravenala, 432-434 Reichelia, 227, 237 Relationships of Metasequoia and Sequoia (Taxodiaceae), The Chromosomes and: an Update, 251-254 Remirea, 226, 233, 237, 240 Renealmia, 16 588 Reproductive Biology of Three Species of Cornus (Cornaceae), Some Observations on the, 419-427 Rhapis excelsa, Anatomy of the Palm, X. Differentiation of Stem Conducting Tis- sue, 191-214 Rhapis, 192, 194, 211 — arborea, 108, 113 — typhina, 411, 416 Rhynchotechum brandisii, 129 237 EK. The Zingiberales (Cannaceae, een ceae, and Zingiber- aceae) in the Southeastern United States, 5-55 Rohria, 228, 237 Romanschulzia, 345 Ronabea, 226, 239 Ropourea, 226, 233, 237 Rorippa, 345, 347, 364, 367, 371 Rosa omeiensis, 61 RosatTt1, THOMAS J. The Plantaginaceae in the eecraiaia United States, 533-562 Rosco Ree 121 — picrodendroides, 121 Rottboelia, 220, 237 Rottboellia, 232, 237 Rourea, 226, 234, 237 Rubiaceae, 238, 240, 534 Rubus giraldianus, 80 — alata, 145, 146 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 65 Rutaceae, 233, 238 Ruyschia, 227, 237 Sabal, 299 Sie 227, 239 Salix cathayana, 61 Salmasia, 227, 237 Salmonia, 230, 237 Salvia (Labiatae) in Mexico, Notes on, with Three New Species, 135-143 Salvia, 135-1 — subg. Calophace, 135 — hispanica, 135 — longiflora, 141, 142 — nervata, | rzedowskii, 139-141 jubifera: 135, 141-143 venosa, 141, 142 Saouari, 224, 227, 237, 239 38 37 Saxifraga micranthidifolia, 61 Saxifragaceae, Schisandra incarnata, 61 — sphenanthera, 61 Schizomeria, 159, 160, 162-165, 167, 168 Schizopetalon, 346 SCHLARBAUM, S. E., T. Tsucuiya, and L. C. JOHNSON. The Chromosomes and Re- lationships of Metasequoia and Sequoia caer an Update, 251-254 Schmidelia, 10 e Clayton, Chemist da Amateur aa 1914- 1978, l- 1984] Schwenkfelda, 227, 237 Scrophulariaceac. 145, 146, 231, 234, 238, 534-536 Seeds of the Cunoniaceae, Fruits and, 149- 19 es 344, 364, 371 a, 37 Set 227, 240 Sequoia (Taxodiaceae), The Chromo- somes and Relationships of Metase- oia and: an Update, 251-254 Boia 251-253 — gigantea, 251 — sempervirens, 251-253 Sequoiadendron giganteum, 251 Sibara, 366, 371 — virginica, 372 Sicelium, 224, 229, 233, 237 Simaba, 217, 227, 237 Simarouba, 120, 227 Simaroubaceae, 109, 120, 238, 240 Simaruba, 234, 237 Simira, 221, 224, 227, 237 Sinapis, 366, 369 — alba, 350, 368 Sinarundinaria indica, 61 Sisymbrium, 347, 367, 373 Sloanea, 216, 217, 219, 232, 237 Smilacina (Liliaceae), Anatomy of Stem Abscission in the Genus, 563-570 Smilacina, 563-570 — henryi, 61 — japonica, 61, 563, 566, 567 — paniculata, 563, 566-569 — racemosa, 61, 563, 564, 566-569 Smithia, 225, 23 7 Solanaceae, 232, 233, 238, 549 Solanum, 217, 237 Some Observations on the Reproductive Biology of Three Species of Cornus (Cornaceae), 419-427 INDEX 589 esate 231 Soramia, 22 Sorbus on 61 Souroubea, 227 Southeastern United States, The Planta- ginaceae in the, 533-562 Southeastern United States, The Tribes of Cruciferae (Brassicaceae) in the, 343-373 Southeastern United States, The Zingiber- ales (Cannaceae, Marantaceae, and Zin- giberaceae) in the, 5-55 Sphagnum, 325 Spigelia, 222, 237 Spiraeanthemum, 150, 155, 164, 165, 167, 169 — katakata, 152, 157, 173 — macgillivrayi, 152, aa — samoense, 152, 1 Spiraeopsis, | Spirematospermum wetzleri, 18 eer y 2 Stanley Seek iece 169, 170 Stauranthera (Gesneriaceae) from New uinea, The First Species of, with Gen- ral Notes on the Genus, 129-1 a co 29 canis winkleri, 129, 131 - cei Songer 129-132 — parvi 2 _ Siilippincasie 129, 130 — pore eee — umbros Stem nes in the Genus Smilacina (Liliaceae), Anatomy of, 563-570 Stem Conducting Tissue, Differentiation of. Anatomy of the Palm Rhapis excelsa, Stemodia, 221, 237 Sterculia, 220, 237 Sterculiaceae, 238 Stollaea, 168 STONE, D. E., C. R. Broome, G. L. 590 WeBsTER, W. J. HAYDEN, and W. T. GiL- Relationship with the Oldfieldioideae (Euphorbiaceae), 105-127 Strelitzia, 432-434 Strelitziaceae, 5, 433, 434, 448 Streptanthus, 344, 347, 348, 366, 369 Streptopus roseus, 61 — streptopoides var. japonicus, 61 Stromanthe, 43, 52, Strychnos, 226, 237 Styrax, 291 Subularia aquatica, 349 Suriana maritima, 113 Systematics and Palynology of Picroden- dron: Further Evidence for Relationship with the Oldfieldioideae (Euphorbiace- ae), 105-127 Systematics of Central American Helico- nia (Heliconiaceae) with Pendent Inflo- rescences, 429-532 Tabebuia ee 416 Tachia Tachi bota, oo a, 170 Selo 218, 219, 228, 237, 239 Tapur: Tarales, 219, 228, 237 Tariri, 228 Tattia, 223, 237 Tauscheria, 349 Taxodiaceae, The Chromosomes and Re- lationships of Metasequoia and Sequoia: an Update, 251-254 Taxodiaceae, 232 Taxodium, 299 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 65 Taxonomic Revision of the American Species of Agarista (Ericaceae), A, 255- 342 Taxonomic Status of Ilex collina and Ne- peices Sear Vegetative my and the, 243-250 fe 344, 364, 369 — nudicaulis, 370 TERABAYASHI, SUSUMU, Davip E. BouF- FORD, and TsUiN-SHEN YING. A Mono- graph of Diphylleia (Berberidaceae), 57- 94 Terebinthaceae, 109 — subg. a a 50 — barbata, 49 _ ieee. 42, 45, 48-50 — divaricata, — geniculata, 45-50, 55 eumoides, 49 Thespesia populnea, 113 Thlaspi, 346, 347, 364, 369 a, 228 Thymocarpus, 40 Tiarella cordifolia subsp. cordifolia, 61 — polyphylla, 61 Tibouchina, 229 R. VINCENT. Anstoniy: of the Palm Rhapis excelsa, X. 1984] Differentiation of Stem Conducting Tis- sue, 191-214 Tonabea, 228, 237 Tonina, 229 ee 229, 237 Tontanea, 224, 229, 237 Tontelea, 229 Topobea, 229 Touchiroa, 216, 229, 237 229 Toxicodendron, 113 — arboreum, 108, 113 Trautvetteria caroliniana, 61 — japonica, Treatment of Aublet’s Generic Names b His Contemporaries and Present-Day Taxonomists, The, 215-242 Tribes of Cruciferae (Brassicaceae) in the Southeastern United States, The, 343- 373 Tn ichocarpus, 216, 237 Tri igonobalanus, a 377, 390, 395, 401 — daichangensis, 376, 377, 381, 389, 390, 394, 395 — excelsa, 376, 377, 391, 395 — verticillata, 376, 377, 383, 390, 394, 395, 40 Trillium erectum, 61 TsucuiyA, T., L. C. JoHNso nd S.E Tummeraceae, 238 Ulmus, 291 Uncaria, 223, 237 Uvaria, 237 — longifolia, 217 INDEX Vaccinium erythrocarpum, 61 aponicum var. japonicum, 61 var. sinensis, 61 vandeina, 221, 237 0, 237 Vegetative Anatomy and the Taxonomic Status of Ilex collina and Nemopanthus (Aquifoliaceae), 243-250 Verbena, 228, 237 ila oe 231, 233, 238 Vero 5 edo 150, 155, 165, 168 — rubifolia, 152, 154-158, 172, 173 Viburnum betulifolium, 61 VINCENT, J. R., and P. B. ToMmLinson. Anatomy of the Palm Rhapis excelsa, X. Differentiation of Stem Conducting Tis- sue, 191-214 Viola acuminata, 61 Violaceae, 239 Virecta, 227 234, 237 Virola, 230 Vochisia, 230, 237 Vochy, 230, 240 Vochysia, 230, 237, 240 Vochysiaceae, 239, 240 Votomita, 230 Vouacapoua, 230, 240 Vouapa, 223, 230, 237, 239 Vouarana, 230 Voyara, 230 Voyria, 230, 239 Waldschmidtia, 216, 237 Warea, 344, 345, 366, 369 _ carter! 370 WesstTerR, G. L., W. J. aa W.T. GIL Lis, D. E. Sto ONE, and C. aren Systematics and a of Picro- dendron: Further Evidence for Relation- ship with the ee (Euphor- biaceae), 105-1 Weinmannia, 150, “satiek 163, 165, 168, 169, 173, 174, 291 — affinis, 152, 156 — aphanoneura, 152 — decora, 153, 156, 157, 174 — denhamii, 153, 156, 158, 174 592 Weinmannia descombesiana, 153, 156, 174 — dichotoma, 153, — fraxinea, 150, 153, 156, 158, 174 — glabra, 153, 173 — hu tchinsonil, 153, 156, 17 — serrata, 153, 237 WHITE, PETER S. The Architecture of Dev- il’s Walking Stick, Aralia spinosa (Ara- areet we 418 Whytocki Willughbeia, oA 237 Willughbeja, 216, 223, 231, 237 Wolfia, 225, 20) Xanthe, 225, 237 JOURNAL OF THE ARNOLD ARBORETUM [voL. 65 Xerodraba pycnophylloides, 349 imenia , 237