4 HARVARD UNIVERSITY VOLUME 60 US ISSN 0004-2625 ao Journal eS the ‘Arnold Arbovetint. : Published: “quarterly in January, Apel; July, and ‘October bi the Arnold Arboretum, Harvard University. Subscription price. $25. 00 per year. eis = oe : X Subscriptions and remittances should be sent to Ms. EB: B. Schmidt, Arnold . ~ Arboretum, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S. 2 cae “ Claims will not be accepted after six months from the date of issue, : Volumes 1-51, ‘reprinted, and some back numbers of volumes 52-56 ‘are available. from the Kraus Reprint Corporation, Route 100, Millwoott ie. New York 10546, U.S.A EDITORIAL COMMITTEE B. G. Schubert, Chairman ad Ore oS Wood, Jr, ASSISTANT EDITOR E. B. Schmidt Printed at the Harvard University pe ores, Boston, ‘Massachusets COVER: The eae, iene, and fruit of Coat caroliniana Walter, hori - beam or blue beech, form the basis of this year’s cover design. An n individual mar was U The designs were all-drawn by Kar n_Stoutsenberger —o Materials a callectsti from plants growing in the ‘Amold Arboretum. ee Widespread in the Northern Hemisphere, Carpinus oo (hiaeatt. in the Betula- ceae or alternatively in the Corylaceae or even the Carpinaceae), with awe. 2 of 35 species, exhibits a disrupted Arcto-Tertiary distribution in eastern America, Europe, Asia Minor, and in eastern Asia where: fc ata of seadick be - diversity occurs. The only North American representativ. le genus, C, .-- caroliniana grows along the edges of streams and in wet, peri forested ¢ so : to - Minnesota, ‘Iowa, Missouri, eastern Texas, and Oklahoma. Disjunct popula- es ie kn own in mountainous regions of southern Mexica and: in Cen tral The stems of Carpinus seantias: a sinall. tree, eS smooth, brownish- or lui uish-gray, sinuate bark and an aj ppear ance reminiscent of those of beech — : trees (Fagus spp.). The wood, which is occasionally used for tool handles, is _ light brown to nearly white. Its growth habit, relatively small. size, conspicuous infructescences, and foliage, which turns deep scarlet and orange to yellow in all, combine to make the species an attractive native ornamental. Sere ee Eas postage paid at Boston, Massachusetts DAVID STURROCK 893-1978 (photograph circa 1946) JOURNAL OF THE ARNOLD ARBORETUM VoL. 60 JANUARY 1979 NUMBER 1 DAVID STURROCK (1893-1978) — MENTOR AND FRIEND RIcHARD A. HOWARD For A PERIOD of ten years, 1936-1946, David Sturrock was superin- tendent of the Atkins Institution of the Arnold Arboretum at Soledad, north of Cienfuegos, Cuba. The garden and collection of tropical plants had been established by Edwin F. Atkins of Belmont, Massachusetts, as the Harvard Botanic Station for Tropical Research and Sugar Cane Research, and in 1932 was formally transferred to Harvard with an en- dowment and a lease of land from the Soledad Sugar Company. It was administered from Cambridge, at first by Dr. Thomas Barbour, Director of the Museum of Comparative Zoology, and later by Dr. Elmer D. Mer- rill, Director of the Arnold Arboretum. Both Barbour and Merrill be- lieved that graduate students in biology at Harvard should have a per- sonal experience in the tropics. To that end, a small grant of money, usu- ally $200 to $300, enabled a thrifty student to travel to Miami or Key West by bus, train, or partway by boat; take a ferry to Havana and a train or bus to Cienfuegos; and spend one to three months in residence at “Soledad.” Room and board charges were modest. Food was Cuban style and consisted of tropical products. The dormitory-type accommoda- tions were at first in Harvard House and later in Casa Catalina. Both buildings had screened porches from which to watch the ever-busy sugar mill and plantations; to overlook the garden and see the sweep of the Trinidad-San Blas Mountains against the horizon; to read, press plants, and work up collections; to enjoy a ‘Sturrock special” before supper or at an evening bridge game; or to take part in the never-ending conversa- tion on the biology of the tropics. The master of this community during my first visit as a student was David Sturrock, a lean, tanned man, usu- ally dressed in white, who spoke Spanish with a Scottish accent — in con- trast to the pure Castilian Spanish of Teodoro, the Spanish-born cook, or the Cuban-dialect Spanish of Manuelo, the houseboy. Sturrock ran a tight ship, making it clear more than once that the money awarded to us for Cuban adventure was money from his budget that might better have been spent on the operations of the garden. We were welcome if we demonstrated a use of the resources assembled, and unwelcome if this © President and Fellows of Harvard College, 1979. 2 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 was only a vacation. Yet it was his influence that directed the research of several students to lifetime investigations in the problems of the An- tilles. David Sturrock was born in Lanark, Scotland, October 28, 1893, one of the eight children of William and Mary Dunn Sturrock. His father, a forester by profession, was the manager of a nursery of fruit and timber trees, with a few ornamentals. Formal schooling under the British system of that time ended at the age of fourteen, and David went to work in a nursery, handling seeds, propagating plants, shipping, planting on contract, pruning, budding, and learning business methods, personnel relations, and plants in the process. At one time he considered an offer from a local businessman to finance his education at Kew Gardens. Local men with more experience advised him against being molded in the Kew fashion, doing only what he was told, and suggested that it would be better instead to learn variety in plants and procedures and to think for himself. He later regarded that as good advice, for further experience in other nurseries broadened his background in the care and use of woody plants. One of his older brothers, who had emigrated from Scotland and now managed a nursery in Miami, offered him employment in the United States. David saved his passage fare plus the $50 reserve then required y the United States Immigration Service and landed in New York in Se ptember, 1913. He was disturbed at the lack of care and appreciation of plants he saw in Central Park and the New York Botanical Garden and without reluctance traveled south to join his brother’s business. His ini- tial assignment in Florida was to locate sources of seeds and scions of ornamental plants useful in landscaping new homes. Without prior knowledge or prejudice, he chose plants from their appearance. Then the company received a contract to landscape the property being developed for the wealthy James Deering. Now known as Viscaya, this land is a current showplace for tourists in the Miami area. Expenditures seemed unlimited, and very large plants were moved with much labor. Many were brought in from the Bahamas, but to locate additional plants Stur- rock was sent to Cuba, where he roamed the island searching for ideal specimens and learning the soil, the natural vegetation, and the farming methods. In the process he met Adolph van Hermann, who owned a small nursery. When the Viscaya project was com pleted, Sturrock resigned from his brother’s firm, moved to Cuba in 1915 on his twenty-second birthday, and accepted the management of a ae s nursery. Van Hermann’s botanical interests were both practical and scientific. He col- lected herbarium specimens which were sent to the New York Botanical Garden, and Sturrock was exposed to the methods of botanical field work. Van Hemiann had a pioneer planting of Mulgoba mango which proved to be Sturrock’s introduction to a plant he was to propagate, breed, and study for the rest of his life. Van Hermann was also concerned with af- forestation of some estates, and Sturrock had the task of developing a timber tree nursery as well. In satisfying his employer’s many interests, Sturrock visited the agricultural experiment station in Santiago de las 1979] HOWARD, DAVID STURROCK, 1893-1978 ) Vegas and met Dr. Juan T. Roig, the prominent Cuban botanist with whom he shared interests for many years. In 1915 David Sturrock, a house guest of the Van Hermanns, was in- vited to join them at a house party given at Christmas by Professor and Mrs. Franklin Sumner Earle in Herradura. Professor Earle had been sent to Cuba by the U. S. Department of Agriculture after the Spanish- American War to establish an agricultural experiment station. He hac also served as the president of the Botanical Society of America and was a specialist in diseases of sugar cane. It was at that party that David Sturrock first met Ruth Earle, whom he married in May, 1918. Their sixtieth wedding anniversary was celebrated six weeks before his death. David Sturrock had applied for United States citizenship in 1913, and within the five-year grace period returned to Miami. The United States was engaged in the first World War, and by enlisting and taking an oath of allegiance Sturrock obtained American citizenship immediately. He served in a medical service unit in France. After the armistice he returned to Cuba, where he accepted a position with the Hershey Sugar Company, reforesting land and developing a nursery for the sale of ornamentals and fruit trees while managing the production of tropical fruits and vegetables that were exported to New York. Mr. Hershey had a sincere interest in the Cuban people and chose David Sturrock to organize a school for orphans from his Central to be trained in agricultural methods. The students had to be at least eight years of age, and by age fifteen were to be discharged from the school and given a posi- tion with the sugar company. Thirty boys were selected for the program, which included experience with crops and animals on the 1,000-acre farm during the mornings, and classes with a resident teacher in the afternoons. The school, known as the Hershey Agricultural School at Aguacate, Cuba, closed for financial reasons in the depression year of 1929. At this point, David and Ruth Sturrock returned to Florida with their three children. David’s older brother and his parents now lived in West Palm Beach, and there David reestablished his nursery and landscaping business as a means of supporting a desire to select and hybridize better varieties of mangoes for Florida. He acquired the Philippine mango from the Earles’ property in Cuba, additional mangoes from the work of Ed- ward Simmonds at the Federal Experiment Station in Miami, and mulgobas from the Van Hermann and Hershey estates. From these stocks David developed an inventory of small plants. Then in 1936 he received a visit from Thomas Barbour, who asked him to succeed the retiring superin- tendent at Soledad, Mr. Robert Grey. The salary was not much, but the opportunity was large and David accepted. His family remained in West Palm Beach during the school year and joined him in Soledad during the summers. Managing the operation at Soledad was a job for an optimist. Its living collections had been decimated by a hurricane the previous year. Its di- rectives were not clear. The budget was prepared and administered through two or more individuals in Cambridge, who visited Soledad on oc- 4 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 casion during the winter months. The donors’ land and activities sur- rounded the garden area and posed problems, especially when oxen, cattle, and horses broke into the garden and nursery. The land available was largely rocky outcrops, unsuitable for growing sugar cane or for pasture, and water sources were inadequate. The Cuban government, under the Batista regime, hardly understood the purposes of the garden but con- trolled, through legislation, the labor and much of the daily operation. Cuba at that time had a one-crop agricultural economic system based on sugar cane. Under Robert Grey the garden’s research effort had been largely in cane. However, the Atkins family had been interested for many years in finding other crops that could aid the economy but that would not compete for labor during the cropping period. Sturrock had a special interest in fruits and timbers, and the garden had the potential of intro- ducing new plants for trial. The seed lists sent to the Arnold Arboretum from botanical gardens throughout the world were used to request seeds, and Sturrock developed a list of seeds from Soledad plants that could be offered in exchange. An inventory of the plants in the garden had been prepared, and under Sturrock’s direction, the total holdings rose to 124 families, 857 genera, and 4396 species. It was certainly the best col- lection of labeled and mapped plants for convenient study anywhere in the American tropics. With the nearby natural vegetation of seacoast, man- grove, serpentine savanna, and mountains, as well as the large Zapata swamp, it was an ideal place in which to teach and learn. Harvard House contained a laboratory and darkroom, a small but adequate library, and a small herbarium based largely on my own collections and those of John George Jack of the Arnold Arboretum. A publication series was started. For a few years a regularly scheduled Harvard class, Biology 215 (Plants of the Tropics), was conducted at Soledad. Visiting professionals were numerous, and Sturrock always did his best to accommodate their needs — from horses for local transport to vials and packaging material for their collections. The major problem at Soledad, however, remained the financial base, never adequate for the potential of the garden and eventually unable to support even restricted operations. During the years of World War IL, the staff, in a patriotic effort, devoted its attention to plants of economic significance in a wartime economy. Two hundred bushels of seeds of Hevea brasiliensis, the rubber tree, were received, germinated, and planted. Four hundred pounds of the lightweight seed of Cryptostegia, the Madagas- car rubber vine, were collected from this milkweed relative and shipped to U.S.D.A. projects in Haiti. One thousand seedlings of Tectona grandis, the teak, were planted. Evaluations were made of fiber sources in the arid land collection called “Utah.” As a result, maintenance of the garden was reduced, and many plants suffered. The war increased the problems of supplies and equipment. When it was decided in Cambridge in 1945 to cut the Soledad labor force in half, Sturrock and Harvard came to a part- ing of ways. Sturrock returned to West Palm Beach but continued his work in Cuba 1979 | HOWARD, DAVID STURROCK, 1893-1978 5 by developing forest plantings for several sugar companies, which he su- pervised until Castro gained control of the island. In Florida Sturrock acquired additional groves of mangoes for further intensive hybridization. Although the fruit was well known, the biology of the plant was not. Con- ventional hand-pollination did not work. Fruit-set was low, and fruit abortion and drop were high. Applications of hormones, successful in other crops, were not successful with mangoes. Seeds were shown to be mono- or polyembryonic, and gametic and nucellar plants developed with different characteristics. Sturrock worked out a method of inarching seed- ling plants by suspending a container-grown seedling within the branches of an adult tree when conventional grafting methods failed. All of these studies resulted in publications by David Sturrock or his son, who chose the mango as subject for his Ph.D. thesis. Eventually David selected and patented two first-generation hybrids. In 1956 a cross of the ‘Edward’ ?,a monoembryonic plant, and ‘Pico’ é , a polyembryonic type, produced the patented ‘Duncan’, a monoembryonic plant, as well as the equally good but unregistered ‘Mekong’ and ‘Wester’, both polyembryonic. Using the monoembryonic ‘Edward’ as the female parent, and ‘Kent’, also mono- embryonic, as the male, Sturrock selected and patented ‘Young’, a mono- embryonic type. Mangoes take between four and seven years to flower and set fruit, and third-generation crosses made in 1966, 1967, and 1968, with the monoembryonic parents ‘Haden’ and ‘Duncan’, produced the ex- cellent ‘Sturrock’ (patent pending) and about thirty other selections still under evaluation. A visit to the Sturrocks’ home during these years of mango testing was a gustatory delight. Guests were asked to examine, test, and evaluate not one but many fresh mangoes in season, and frozen pulp at other times. His interest in tropical fruits was wide. While at Soledad, and later in West Palm Beach, David and his wife grew and tested hundreds of tropical fruits, working out methods of preparation and preservation. The first publication of the Atkins Institute of the Arnold Arboretum was Tropical Fruits for Southern Florida and Cuba and Their Uses. This be- came known as “the jelly book” and is widely quoted today in the litera- ture of tropical botany. Recipes prepared by the Sturrocks appeared in most of his writings, and a later publication, Fruits for Southern Florida, appeared serially in the Stwart News before publication as a hardcover ok. This has been reprinted once and is scheduled for an additional printing in 1979. His favorites, as I recall with pleasure, were a sherbet made of Carissa macrocarpa, a jelly of Carissa karandas, and, for eating out of hand (second only to mangoes), Myrciaria floribunda (jaboticaba). In cooperation with Dr. Edwin A. Menninger, Sturrock compiled his notes on tropical ornamental plants. The volume Shade and Ornamental Trees for South Florida and Cuba was originally published in serial form in the Stuart News. This was the first volume for Florida that paid at- tention to salt-resistant trees for exposed coastal sites, the areas of deep, overdrained sands, the shallow areas over rock, and the poorly drained areas inland. It is one of the few volumes in tropical and subtropical hor- 6 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 60 ticulture that emphasize the ecology of the planting site. Again, his wife contributed to his work, this time by supplying the drawings of the plants. Sturrock’s work with potential timber trees climaxed in perhaps his least-appreciated volume, Trees for Southern Florida: A Plea for Affores- tation, which was published by the Central and Southern Florida Flood Control District in West Palm Beach. Throughout his life, in Scotland, Cuba, and Florida, Sturrock saw lands denuded of trees needed for timber, windbreaks, or shade. His experience in Soledad gave him knowledge of the species with the most rapid growth and the desirable shape or root system. His reports to his superiors at Harvard are dominated by pages of notes, observations, and recommendations on timber trees at Soledad that were never included in published annual reports. He was deeply shocked by the report that his growth study plantations at Soledad and his timber plantings elsewhere in Cuba had been cut quickly and completely for charcoal in the early stages of the Castro administration. Late in his life, David Sturrock turned to wood carving for relaxation. He experimented with a variety of tropical woods to achieve the minute detail he sought, at first carving scenes from Cuba which his wife sketched in outline. His work was accepted artistically and scientifically, and fif- teen carved panels, representing Florida trees and Seminole Indians. are displayed in the public library of West Palm Beach. Thirteen panels, rep- resenting the vegetation of different geologic periods, and a twenty-foot panel on the evolution of man, are in the Science Museum and Planetarium of Palm Beach County. David Sturrock preferred to be called a “plantsman” for his broad in- terest in all plants and their environment. He indeed had a comprehensive knowledge and many talents. To the student he was patient, understand- ing, and willing to share any knowledge he had. To me he was mentor and friend. He died July 8, 1978, and was buried in Hillcrest Memorial Park in West Palm Beach, Florida. He is survived by his wife, Ruth Esther Earle Sturrock, and four children, of whom he was justly proud: James P. Sturrock, an aerospace engineer; Thomas T. Sturrock, a profes- sor of botany and administrator; Ruth Mary Sturrock, an electron micro- scopist; and Peter E. Sturrock, a professor of chemistry. PUBLISHED WRITINGS OF DAVID STURROCK 1939. Rootstock experiments with Malpighia. Proc. Florida State Hort. Soc. 52: 42, 43 1940. Tropical fruits for southern Florida and Cuba and their uses. Publ. Atkins Inst. Arnold Arb, 1: 1-131. 1944. Notes on the mango. 122 pp. Stuart Daily News, Inc., Stuart, Florida, published in cooperation with the Atkins Institution of the Amold Arboretum. 1946. Shade and ornamental trees for South Florida and Cuba. (With E. A. Menninger.) 172 pp. Stuart Daily News, Inc., Stuart, Florida. 1947, The influence of the Indian and Philippine mangos on the development of the mango in Florida. Proc. Florida Mango Forum 1947: 20-24. HOWARD, DAVID STURROCK, 1893-1978 7 The pruning of mango trees. /bid. 1948: 40-44. Karanda as a commercial fruit. Proc. Florida State Hort. Soc. 61: 289- 1. Frost protection for young mango trees. Proc. Florida Mango Forum 1949: Wrapping oe mango trees for frost protection. /bid. 1951: 13-16. Stem protection of young fruit trees from frost. Proc. Florida State Hort. Soc. 64: 262-264 Notes on vitamin content of mangos. Proc. Florida Mango Forum 1952: 40-46. Soe affecting commercial mango ere eas in Martin County. Proc, Florida State Hort. Soc. 72: 320-32 Fruits for Southern Florida. 196 pp. eee Printing Co., Stuart, Florida Progress report on some mango hybrids. Proc. Florida State Hort. Soc. 72: 384-387 Trees for southern Florida: A plea for afforestation. 242 pp. Central & Southern Florida Flood Control District, West Palm Beach, Florida. Final report on some mango hybrids — 1969. Proc. Florida State Hort. oc. 82: 318-321 ARNOLD ARBORETUM OF HARVARD UNIVERSITY 22 DIVINITY AVENUE CAMBRIDGE, MASSACHUSETTS 02138 8 JOURNAL OF THE ARNOLD ARBORETUM [| VoL. 60 LEAF ANATOMY AND VENATION PATTERNS OF THE STYRACACEAE } WILLIAM E. SCHADEL AND WILLIAM C. DICKISON THE STYRACACEAE is a woody, dicotyledonous family that is generally described as containing 12 or 13 genera and 150 to 190 species (Wood & Channell, 1960; Hutchinson, 1973; Gonsoulin, 1974; Spongberg, 1976). The plants are distributed primarily in eastern Asia to western Malaysia, tropical South America, and the southeastern United States. Ecologically, the plants occupy a variety of habitats. The largest genus, Styrax L., includes about 120 species and is wide- ly distributed in the tropical and warm temperate regions of eastern Asia and America. A conspicuous exception to this distribution pattern is Styrax officinalis L., which is found in the Mediterranean region and in California. In Asia the numerous species extend from the eastern Hima- layas to Malaysia, Indochina, Korea, and Japan. The New World species are widely distributed from Pacific to Atlantic North America, the West Indies, southward to Venezuela and southern Brazil. Two sections of the genus were recognized by Perkins (1907): STYRAXx (Eustyrax Perk.), with the ovary 16- to 24-ovulate, and FovrEoLartA (Ruiz & Pavon) Perk., with the ovary 3- to 5-ovulate. Section StyRax was divided by Giirke (1891) and Perkins (1907) into two series, Valvatae and Imbricatae, on the basis of aestivation of the corolla lobes. As van Steenis (1932) has shown, however, imbrication varies widely with re- spect to both individual plants and species. The genus Halesia Ellis ex Linnaeus (about 5 species) occurs in southeastern North America and in eastern Asia, Bruinsmia Boerlage & Koorders (2 species) is found in As- sam, Burma, and Malaysia (excluding the Malay Peninsula), Pamphilia Martius ex A. de Candolle (3 species) is located in Brazil, and A frostyrax Perkins & Gilg (2 species), which according to Baas (1972) is better placed in the family Huaceae of the Malvales, occurs in tropical Africa. The remainder of the genera are distributed from the Himalayas through China and Japan: Alniphyllum Matsumura (8 species), China and For- mosa; Pterostyrax Siebold & Zuccarini (7 species), Burma to Japan; Reh- derodendron Hu (10 species), China and Indochina; Huodendron Rehder (6 species), southern China, Siam, and Indochina; Sinojackia Hu (3 spe- cies), southern China; Parastyrax W. W. Smith (1 species), Burma; and Melliodendron Handel- Mazzetti (3 species), southern and southwestern China. As reviewed by Spongberg (1976), the flowers of the Styracaceae are * This study eee a portion of a thesis submitted by W. E. Schadel to the Graduate School, The University of North Carolina at Chapel Hill, in partial fulfill- ment of the Larne for the Master of Arts degree 1979 | SCHADEL & DICKISON, LEAF ANATOMY 9 perfect and actinomorphic. Inflorescences are often reduced, and the flowers are usually arranged in simple or sometimes branched, terminal and/or axillary racemes, corymbs, or panicles and are sometimes solitary or in few-flowered fascicles. The calyx is often articulated at the base, synsepalous, tubular, subentire or 4- or 5-lobed (rarely more), often united into a tube only at the base. The insertion of the corolla is hypogynous to epigynous, and the lobes are valvate or imbricate in bud. The stamens are usually twice the number of the corolla lobes, sometimes more; epipe- talous; inserted in 1 whorl on the base of the corolla (or rarely at the base of the ovary), with the filaments usually being connate into a tube below (free above) and the anthers being 4-locular. The gynoecium is syncarpous. The ovary is superior and completely free to inferior and completely adnate to the calyx tube and is 3- to 5-locular, often 1 -locular above. Each locule has 1 to many ovules on axile placentae. Each flower has a single slender style terminated by a usually 3- to 5-lobed stigma. The fruits are dry, often woody, dehiscent or indehiscent capsules or drupes, and the seeds number from 1 to many. The embryo is usually straight with broad cotyledons and is embedded in a fleshy endosperm. In the general systems of Bentham and Hooker (1876), Engler and Prantl (Perkins, 1907), Bessey (1915), Cronquist (1968), Thorne (1968, 1976), and Takhtajan (1969, 1973), there is general agreement on the placement of the Styracaceae in the Ebenales. This same treatment was followed by G. Wagenitz (1964) in the latest edition of Engler’s Syllabus der Pflanzenfamilien. Hutchinson (1926, 1959, 1967, 1969, 1973), however, placed the family in the order Styracales, along with the families Lisso- carpaceae and Symplocaceae. The families Ebenaceae, Sapotaceae, and Sarcospermataceae were considered to constitute the distinct order Ebenales. Hutchinson (of. cit.) derived the Styracales from the Rosales via the Cu- noniales and derived the Ebenales (sensu stricto) from the Myrsinales. Huber (1963) suggested a relationship between Symplocaceae and Styraca- ceae and the Cornaceae, incorporating them in the order Cornales which he derived from the Cistiflorae, especially Theaceae. Airy Shaw (in Wil- lis, 1973) denied affinities between Symplocaceae and Styracaceae. Ac- cording to Airy Shaw (op. cit.), Styracaceae is related to Philadelphaceae (also placed in the Cornales by Huber), and Symplocaceae is related to Theaceae. Cronquist (1968), Thorne (1968, 1976), and Takhtajan (1969, 1973) were of the opinion that the Ebenales, including the families Sapota- ceae, Ebenaceae, Symplocaceae, Styracaceae, and Lissocarpaceae, form a natural group and that the origin of the order lies among the thealean complex. According to Wood and Channell (1960), these 5 families are characterized by sympetalous corollas with stamens generally two to three times as many as the corolla lobes (or, by abortion, equal in number to and opposite them) and by superior to inferior, incompletely to completely loculed ovaries with axile placentation. Wagenitz (1964) concluded that the Ebenales should also include two additional families, Hoplestigmata- ceae and Sarcospermataceae. Dahlgren (1975) felt that only the families Styracaceae, Sapotaceae, Lissocarpaceae, and Ebenaceae comprised the 10 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 Ebenales and that the Symplocaceae was better placed in Theales. Ac- cording to Nooteboom (1975), Symplocaceae does not belong in the Ebe- nales but shows more affinity to Cornaceae and possibly Theaceae on the basis of evidence derived from chromosome numbers, chemotaxonomy, anatomy (wood and leaf), germination studies, and a survey of morpho- logical characters. Such arguments indicate that the order Ebenales (sensu lato), including the families Lissocarpaceae, Styracaceae, Ebenaceae, Sapo- taceae, and Symplocaceae, may not be a very natural assemblage and is in need of redefinition. The first comprehensive anatomical investigation of styracaceous leaves was the family treatment by Solereder (1908). Metcalfe and Chalk (1950) provided additional observations, but information regarding the leaf anat- omy of this family is still very incomplete. Other literature containing general information about leaf characters includes work by Giirke (1891), Perkins (1907, 1928), Mori (1936), Mears (1972), Baas (1972), Fahn (1974), Gonsoulin (1974), Hickey and Wolfe (1975), and Sarcar and Banerji (1975). The present research was undertaken to examine the leaf anatomy (in- cluding venation patterns) of the Styracaceae in order to provide additional evidence for generic circumscription and to shed additional light on the intrafamilial and interfamilial relationships of the family. The specific objectives of this initial investigation are: 1) to provide a comprehensive description of the leaf structure of the Styracaceae, 2) to outline basic trends of structural specialization in the anatomy and architecture of styracaceous leaves, and 3) to provide anatomical data that will eventually contribute to a better understanding of the intrafamilial and interfamilial systematics and relationships of the family. MATERIALS AND METHODS Leaves from all 12 genera recognized as comprising the family Styraca- ceae (sensu Hutchinson, 1973) were studied. Material of the related genus Lissocarpa (Lissocarpaceae, 2 species, tropical South America) was also examined. A total of 82 species were investigated (APPENDIX). Trans- verse sections of leaves were prepared according to standard procedures of paraffin embedding and sectioning (Johansen, 1940). Sections were taken from the central part of the lamina (including the midrib) and from the basal, medial, and distal portions of the petiole. Since almost all specimens were gathered from dried herbarium collections, it was necessary to reexpand the leaves initially in 5 percent NaOH before fixation in 70 percent FAA and dehydration. Transverse sections were stained with safranin and fast-green. Cleared leaves were prepared using the NaOH method outlined by Arnott (1959) and were subsequently stained with safranin. Using cleared leaves and temporary free-hand peels, stomatal patterns were observed. Ten stomata per specimen were measured at the widest part of the guard cell pair and from pole to pole. Averages and ranges for stomatal width and length were recorded. Drawings were made 1979 | SCHADEL & DICKISON, LEAF ANATOMY aft using a Wild dissecting microscope with a drawing tube attachment. Pho- tomicrographs of transverse sections and cleared preparations were made using a Zeiss photomicroscope. For the description of leaf architecture, the terminology of Hickey (1973), with necessary additions and modifications, was followed. For the genus Styrax the masculine gender was implemented following the sug- gestion of Nicolson and Steyskal (1976). Slides are deposited in the Department of Botany, The University of North Carolina at Chapel Hill, and in the Department of Botany, Smith- sonian Institution, Washington, D. C. OBSERVATIONS SUMMARY OF LEAF ANATOMY OF THE STYRACACEAE (sensu stricto) Petiole transverse section. Cuticle smooth or minutely furrowed. Epi- dermis uniseriate. Epidermal cells irregular to more or less rectangular to square in outline and with moderately thin walls. Parenchymatous cor- tex undifferentiated except in the monotypic genus Parastyrax, where outer half is collenchymatous and inner half parenchymatous. Crystals occurring as scattered druses, scattered prismatic crystals, a combination of druse and prismatic crystals, or, as in Parastyrax, crystal sand. Nu- merous solitary and clustered brachysclereids present in Huodendron. Vasculature of petiole variable within the family. Five different vascu- lar configurations occurring in basal portion of petiole: 1) arc, 2) arc with two dorsal cortical bundles, 3) arc with invaginated ends, 4) arc with invaginated ends and two dorsal collateral bundles, and 5) dissected cylinder of collateral bundles with numerous medullary bundles and four smaller dorsal dissected cylinders. At distal end of petiole, the following patterns occur: 1) Arc with invaginated ends: Styrax americanus var. americanus (Prate III, C), S. americanus var. pulverulentus, S. hypargyreus, Oe paralleloneurus. bo — Arc with invaginated ends accompanied by two dorsal cortical bundles: Pamphilia aurea (Pirate III, A), P. pedicellata var. ovalis, P. styracifolia, Styrax argenteus var. argenteus, S. argenteus var. lintonii, S. argenteus var. ramirezii, S. benzoin (PLaTE IV, A), S. camporum, S. formosanus, S. glaber, S. glabrescens var. glabrescens, S. glabrescens var. pilosus, S. grandifolius, S. henryi, S. hypoglaucus, S. japonicus (Pate III, D), S. langkongensis, S. obtusifolius, S. officinalis var. fulvescens, S. officinalis var. redivivus, S. philadel- phoides, S. platanifolius, S. portoricensis, S. subcrenatus, S. subert- folius, S. sumatranus. Ww eae Medullated cylinder: Halesia macgregorii, Huodendron biaristatum, H. chunianum, H. tibeticum, Sinojackia henryi, S. rehderiana, S. xylocarpa (PLATE III, B), Styrax agrestis, S. faberi. 12 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 4) Medullated cylinder accompanied by two dorsal cortical bundles: Al- niphyllum eberhardtii, A. fortunei, A. hainanense, A. pteros pe Halesia carolina, H. diptera var. diptera, H. diptera var. ee pe Melliodendron wangianum, M. xylocarpum, Pterostyrax cavaleriei, P. corymbosus, P. hispidus, Rehderodendron indochinense, R. kwei- chowense (PLATE II, B), R. macrocarpum, Styrax ochraceus (PLATE IV, C), S. serrulatus. Sal — Medullated cylinder accompanied by medullary vascular tissue and two dorsal cortical bundles: Bruinsmia styracoides, Styrax am- biguus, S. cespedesii, S. conterminus, S. crotonoides, S. davillifolius, S. fanshawei, S. ferrugineus, S. glabratus, S. heteroclitus, S. jaliscanus, S. latifolius, S. longifolius, S. macrocalyx, S. macrotrichus, S. martii, S. micrasterus (PLATE IV, B), S. microphyllus, S. ovatus, S. pallidus, S. pearcei, S. pentlandianus, S. pohlii (Prater IV, D), S. poisson- ianus, S. pseudargyrophyllus, S. subcordatus, S. yapobodensis. ON — Dissected cylinder of collateral bundles with numerous medullary bundles and two smaller dorsal dissected cylinders: Parastyrax lacei (Prats II, 3A.) All cortical bundles in the family collateral except in Styrax argenteus, S. davillifolius, and S. pohlii (PLATE IV, D), where amphicribral bundles occur. Perivascular sclerenchymatous elements present distally. As vas- cular tissue enters lamina, configuration remains same as in distal portion of petiole for all species investigated except Pamphilia (PLATE ITI, A) and Parastyrax (PLATE IT, A). Petiole and lamina surface view. Trichomes present in all species in- vestigated except Halesia macgregorii. They generally occur on petiole, as well as on both surfaces of lamina, in four combinations of structural types: 1) stellate only, 2) peltate only, 3) stellate and peltate, and 4) stellate and simple cylindrical. Cuticle smooth or striated. Within indi- vidual cells striations oriented in a single direction parallel to one another and sometimes continuous from cell to cell. Laminar epidermal cells of both surfaces varying in shape from irregular to more or less quadrangular to polygonal. Epidermal anticlinal cell walls moderately thin except in Styrax argenteus, S. camporum, S. heteroclitus, and S. macrotrichus, where moderately thick walls occur. Anticlinal cell walls of adaxial epidermis generally straight to slightly curved, being slightly sinuous to curved only in Huodendron biaristatum, H. chunianum, Styrax glabrescens, and S. ser- rulatus. Anticlinal cell walls of abaxial epidermis straight to slightly curved, slightly sinuous to curved, or sinuous. For all species investigated, stomata anomocytic, occurring only on abaxial surface, and ranging in mean width between 11 and 22 ym. and in mean length between 14 and 80 pm). Venation pinnate, with the single midvein following a straight, un- branched course. Midvein of moderate size (1.25-2.00 percent) except 1979] SCHADEL & DICKISON, LEAF ANATOMY 1) in Sinojackia (PLATE VI, E) and Styrax platanifolius, where it is thin (less than 1.25 percent). Midvein ensheathed by sclerenchymatous ele- ments elongate along long axis. In Parastyrax (Prate VI, B) venation actinodromous with a pair of lateral primary veins occurring on either side of a third, centrally located primary vein. The imperfectly developed, reticulate actinodromous lateral veins becoming attenuate exmedially and with secondary veins originating exmedially. In all pinnately veined species, secondary venation weakly cladodromous, eucamptodromous, or transitional from eucamptodromous basally to brochidodromous distally. In species that are brochidodromous distally, loop-forming branches of secondaries join superjacent secondaries at right and acute angles. Rela- tive thickness of secondary veins moderate, and their course branched when cladodromous, uniformly curved when eucamptodromous, and abrupt- ly curved when brochidodromous. Exmedially, they are attenuate except for brochidodromous veins, which are of uniform thickness, and are braced in all species by a system of tertiary and quaternary veins. Sim- ple intersecondary veins occurring in all species investigated except Halesia macgregorii (PLATE V, G), Rehderodendron (PiaTE VI, D), Sinojackia (Pirate VI, E), Styrax conterminus, S. martii, S. officinalis (PLATE VI. I), S. subcordatus, S. subcrenatus, S. suberifolius, and S. yapobodensis, in which intersecondary veins are composite. Intramarginal veins absent in all species. Tertiary veins originating from secondary veins at acute, right, and obtuse angles, their pattern varying within the family from percur- rent to random reticulate. Percurrent tertiary veins simple to forked and close in all species in which they occur, except in the specimen of Para- styrax examined, where they are both closely and distantly arranged. Pre- dominantly opposite percurrent tertiary veins have a constant oblique tertiary angle in Parastyrax, Styrax argenteus (Pate VI, G), and Styrax benzoin, and an apically decreasing oblique tertiary angle in Alniphyllum eberhardtii (PLATE V, C), Pterostyrax (Piate VI, C), and Styrax jali- scanus, Predominantly alternate percurrent tertiary veins have a constant oblique tertiary angle with the midvein in Styrax cespedesti and Styrax macrotrichus, and an apically decreasing oblique tertiary angle in Alni- phyllum fortunei, A. hainanense, A. pterospermum (PLATE V, D), Stvrax crotonoides, S. heteroclitus, and S. latifolius. In other species, tertiary veins percurrent only in basal portion of leaf and forming random reticu- late patterns distally. Predominantly opposite percurrent tertiary veins occurring only in a basal position in Huodendron chunianum, Styrax con- terminus, S. ovatus, S. paralleloneurus, and S. yapobodensis; predominant- ly alternate percurrent tertiary veins, only in a basal position in Mellio- dendron (Pate V, I), S. ambiguus, S. ferrugineus, S. glabrescens, S. hen- ryi, S. hypoglaucus, S. langkongensis, S. longifolius, S. macrocalyx, S. mar- tii, S. pentlandianus, S. poissonianus, S. pseudargyrophyllus, S. serrulatus, S. subcordatus, S. suberifolius, and S. sumatranus. In all other cases, tertiaries form random reticulate patterns throughout leaf. Resolution of higher vein orders distinct. Quaternary and quinary veins follow a rela- tively random course, having bundle sheaths of either parenchymatous or 14 JOURNAL OF THE ARNOLD ARBORETUM [voL. 60 sclerenchymatous elements elongate along their long axes. Bundle sheaths predominantly sclerenchymatous, with parenchymatous sheathing occur- ring only in Alniphyllum, Bruinsmia, Parastyrax, Styrax americanus, S. argenteus, and S. platanifolius. Ultimate veinlets simple to branched, lin- ear to curved, generally with weak bundle sheaths composed of elements similar to sheathing of their respective higher vein orders. Sheathing ab- sent on ultimate veinlets in Alniphyllum, Bruinsmia, Halesia (except H. macgregorit, where present), Melliodendron, Parastyrax, and Pterostyrax. Marginal ultimate venation either incomplete or looped. Areoles irregular to quadrangular to polygonal in shape (PLATE VII, A-F). They are small (less than 0.3 mm.) in Styvax crotonoides, S. pseudargyrophyllus, and S. suberifolius; small and medium (0.3-1.0 mm.) in Parastyrax, Styrax paralleloneurus, and S. sumatranus; large (1.0-2.0 mm.) in Styrax ja- ponicus; both medium and large in Bruinsmia: and medium in all other species investigated. Areolation imperfectly or well developed, being in- complete only in the specimens of Styrax japonicus examined. Areoles relatively randomly arranged except in Huodendron biaristatum, H. chu- nianum, Sinojackia, Styrax crotonoides, S. pseudargyrophyllus, S. sub- crenatus, and S. suberifolius, where well developed and relatively highly oriented. Minute, crenulate teeth present in some species investigated are vascu- larized by a tertiary vein that is slightly expanded at base of tooth and tapered near apex (PLATE VIII, A-D). A glandlike cap composed of thin- walled cells occurring at apex of each tooth. Such glandlike structures also occurring on the entire margins of Styrax glabrescens. Scattered cir- cular glands, consisting of five to eight small, flattened cells surrounding a centrally depressed area, occurring on lower leaf surface of a few species of Styrax. Lamina transverse section. Cuticle thicker on adaxial surface than on abaxial in about half of species investigated, and of equal thickness on both surfaces for other species. For all species adaxial cuticular thickness ranging from less than 1 ym. to 24 um.; abaxial cuticular thickness, from less than 1 um. to 4 ym. Epidermal cells of both surfaces thin -walled and irregular to more or less rectangular to a Epidermis uniseriate ex- cept in Styrax macrotrichus (PLATE X, A), S. micrasterus (PLATE X, B), S. portoricensis (PLATE X, E), and S. pseudargyrophyllus (PLATE X, F), where biseriate epidermis (hypodermis?) is present. Abaxial epidermal cells smaller than adaxial cells. Anomocytic stomata level with or shightly raised above abaxial surface. All major veins raised above abaxial leaf surface. Bifacial mesophyll composed of 1 or 2 layers of slightly to very elongate palisade parenchyma cells in all species except Styrax ochraceus (PLATE X, C), which has 2 or 3 distinct layers of palisade cells, and Styrax macro- trichus (PLATE X, A), S. poissonianus, and S. portoricensis (PLATE X, i), with only a weakly differentiated mesophyll of irregular to more or less rectangular parenchyma cells. Spongy tissue very compact, moderately 1979] SCHADEL & DICKISON, LEAF ANATOMY 1S compact, or loose, generally consisting of more or less isodiametric to slightly lobed parenchyma cells. Secondary and tertiary veins with scler- enchymatous bundle sheaths composed of elements elongate along their long axes. Bundle sheath extensions present in some species, occurring in two patterns of positions, adaxial only, and adaxial and abaxial. They are composed of sclerenchymatous elements elongate along long axis of veins. In Pamphilia aurea Martius (Plate IX, D) (/rwin et al. 20974 and 29208) some elements of bundle sheath extensions also elongate perpendi- cular to long axis of veins. Quaternary and quinary veins with pre- dominantly sclerenchymatous sheaths (occurring only in Alniphyllum, Bruinsmia, Parastyrax, Styrax americanus, S. argenteus, and S. platani- folius) and generally lacking bundle sheath extensions. Crystals present in all plants investigated, occurring as scattered druses, scattered prismatic crystals, a combination of druse and prismatic crystals, or, as in Para- styrax, crystal sand. They are sometimes confined to cortical parenchyma of midrib and to parenchyma cells adjacent to veins. Vasculature of mid- rib variable within the family, consisting of an arc, usually with invaginated ends, a medullated cylinder, or a medullated cylinder with medullary vas- cular tissue. Midrib with perivascular sclerenchymatous elements in all species investigated except Styrax americanus. In Parastyrax the central actinodromous vein has a ring of collateral bundles surrounding a single collateral bundle. All collateral bundles have thin-walled sclerenchymatous caps. Lissocarpa Bentham (Lissocarpaceae, sensu Engler & Gilg, 1924) Petiole transverse section. Cuticle smooth, epidermis uniseriate. Epi- dermal cells irregular in outline and with moderately thick walls. Paren- chymatous cortex undifferentiated, containing prismatic crystals and nu- merous diffuse vesiculose sclereids (sersu Rao & Bhupal, 1973) irregular in outline. Basally, vascular tissue consists of an arc with invaginated ends accompanied by two dorsal collateral bundles (PLATE I, B). Medially, each invaginated end of the arc begins to fold back upon itself dorsally. Distally, dorsal folding of each invaginated end of the arc increases. Peri- vascular sclerenchymatous elements present both singly and in clusters. As vascular tissue of petiole enters lamina, products of dorsal folding separate from arc and subsequently fuse, forming a dorsal plate. Ends of arc also fuse, forming a medullated vascular cylinder. Petiole and lamina surface view. Surface of petiole and lamina glab- rous. Cuticle smooth. Adaxial epidermal cells of lamina irregularly shaped with moderately thick, extremely sinuous anticlinal walls. Abaxial epi- dermal cells resembling cells of upper surface. Diffuse vesiculose sclereids with numerous simple pits occurring singly and in clusters throughout meso- phyll and in random association with vascular tissue. For the genus, sto- mata anomocytic, occurring only on abaxial surface, ranging in mean width between 28 and 29 »m., and in mean length between 34 and 38 pm. 16 JOURNAL OF THE ARNOLD ARBORETUM [voL. 60 Venation pinnate, with single midvein following a straight, unbranched course. Midvein of moderate size (1.25—2.00 percent) and ensheathed by thick-walled sclerenchymatous elements. Secondary venation weakly bro- chidodromous and ensheathed by thick-walled sclerenchymatous elements ee along long axis of veins. Acute angle of divergence, nearly uni- form for all re is wide. oe veins relatively slender, their course abruptly curved: loop-forming branches of secondaries joining su- perjacent secondaries at right and acute angles and braced exmedially by system of tertiary and quaternary veins. Composite intersecondary veins present, but intramarginal veins absent. Tertiary veins originate from secondaries at acute, right, and obtuse angles, forming random reticulate pattern. They are ensheathed by moderately thick-walled sclerenchyma- tous elements elongate along long axis of veins. Resolution of higher vein orders not distinct. They consist of a ramified network terminating in nu- merous branched ultimate veinlets. Higher vein orders and ultimate vein- lets ensheathed by thin-walled, irregular to rectangular sclerenchymatous elements elongate along their long axes. Marginal ultimate venation fim- briate. Imperfectly developed, randomly arranged areoles irregularly shaped and large (1.0-2.0 mm.). Teeth and glands absent. Lamina transverse section. Cuticle thicker adaxially (5-6 »m.) than abaxially (3-4 um.). Epidermis uniseriate. Epidermal cells of adaxial surface thin-walled and irregular to more or less rectangular in outline; those of both surfaces approximately equal in size. Anomocytic stomata level with abaxial leaf surface. All major veins raised above abaxial lea surface. Bifacial mesophyll with 1 or 2 layers of short, more or less rectangular palisade parenchyma cells and several layers of fairly loose spongy tissue of isodiametric to slightly lobed cells. Sclerenchymatous bundle sheaths without bundle sheath extensions are conspicuous around all orders of vena- tion. Prismatic crystals confined to parenchyma cells adjacent to veins. Diffuse vesiculose sclereids, irregularly shaped in outline, occurring in mesophyll. Midrib with medullated vascular cylinder accompanied by dorsal plate. Perivascular sclerenchymatous elements present. Afrostyrax Perkins & Gilg Petiole transverse section. Cuticle smooth, epidermis uniseriate. Epi- dermal cells square to rectangular in outline and with moderately thin walls. Parenchymatous cortex undifferentiated except in one specimen of Afrostyrax kamerunensis (Mildbraed 10688 (A)), where small amount pean of periderm and 2 or 3 layers of collenchyma observed in outer portion of cortex. Cortex containing prismatic crystals. Basally and medially, vascular tissue of petiole consisting of two large, C-shaped vascular bun- dles and one smaller, abaxially positioned one (Pirate I, A). Distally, the two large, C-shaped lateral bundles invaginate at the dorsal ends. Peri- vascular sclerenchymatous elements present. As vascular tissue of petiole 1979] SCHADEL & DICKISON, LEAF ANATOMY 17 enters lamina, invaginated dorsal ends of C-shaped lateral bundles sep- arate from lateral bundles, subsequently becoming medullary bundles. The two lateral C-shaped bundles fuse dorsally, enclosing the medullary bun- dles, which vary in number from one to several depending on whether in- vaginated portions fuse or divide. Baas (1972) reports A. ee ome as having a more complex vascularization than A. kRamerunensts as the r sult of a larger, although variable, number of medullary bundles see Petiole and lamina surface view. Stellate trichomes occurring infre- quently on petiole and lamina of both species. Numerous peltate scales and short, uniseriate, two- to three-cellular trichomes occurring on petiole and lamina surfaces of A frostyrax lepidophyllus but lacking on A. kamer- unensis. Cuticle smooth. A frostyrax lepidophyllus with adaxial epidermal cells of lamina quadrangular to polygonal, and having thin, slightly sin- uous to curved anticlinal walls; A. Ramerunensis, however, with anticlinal epidermal walls very sinuous. With exception of cells underlying major veins and resembling cells of adaxial surface, abaxial epidermis of 4. lepidophyllus almost completely composed of dome-shaped papillate cells. This condition absent in A. kamerunensis, where abaxial epidermal cells smaller but otherwise identical to upper surface cells (which have very sinuous anticlinal walls). For the genus, stomata paracytic with crescent- shaped subsidiary cells, occurring only on abaxial surface. Mean stomatal width 11 um.; mean length ranging between 15 and 16 um. Venation pinnate, with single midvein following a straight, unbranched course (PLATE V, A). Midvein of moderate size (1.25—2.00 percent) en- sheathed by moderately thick-walled sclerenchymatous elements elongate along long axis of midvein. Secondary venation transitional from eucamp- todromous basally to brochidodromous distally, also ensheathed by mod- erately thick-walled sclerenchymatous elements elongate along long axis of veins. Acute angle of divergence, nearly uniform for all secondaries, moderate. Relative thickness of the secondary veins moderate, their course proximately right angles and forming orthogonal reticulate patterns with other tertiaries. They have a sheath of thin-walled parenchymatous cells slightly elongate along long axis of veins. Resolution of higher vein orders distinct. Both quaternary and quinary veins following an orthogonal course and having weak, incomplete parenchymatous sheaths. The few ultimate veinlets simple, linear to curved, without sheathing. Marginal ultimate venation fimbriate. The well-developed, oriented areoles quad- rangular and small (less than 0.3 mm.) (PLaTE VII, A). Teeth absent, but oval-shaped glands composed of elongate, thin-walled cells present on lamina margins. Lamina transverse section. Cuticle smooth and of equal thickness (less than 1 um.) on both surfaces. Epidermis uniseriate. Epidermal cells of adaxial surface thin-walled and square to rectangular in outline. 18 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 Cells of abaxial surface smaller than adaxial cells. Majority of abaxial cells of Afrostyrax lepidophyllus papillate with an outer dome-shaped con- figuration (PLATE IX, A), a condition absent in A. kamerunensis. Para- cytic stomata slightly raised above level of abaxial leaf surface. All major veins raised above level of abaxial leaf surface. Bifacial mesophyll with 1 or 2 layers of elongate palisade parenchyma cells and two to several layers of moderately compact spongy tissue of more or less isodiametric to slightly lobed cells. Sclerenchymatous ele- ments ensheathing secondary veins and forming adaxial and abaxial caps exterior to parenchymatous bundle sheaths of tertiaries. Both secondary and tertiary veins with conspicuous parenchymatous adaxial and ee bundle sheath extensions. Quaternary and quinary veins with weak, complete parenchymatous sheaths and parenchymatous adaxial and ea bundle sheath extensions. Prismatic crystals occurring in parenchyma cells adjacent to veins and in cortical region of midrib. Midrib with medul- lated vascular cylinder having variable number of medullary bundles. Perivascular sclerenchymatous elements present. DISCUSSION AND CONCLUSIONS Leaves of Styracaceae are considerably more variable morphologically and anatomically than previous descriptions indicate. Characters that are present in leaves of nearly all species investigated include bifacial meso- phyll, anomocytic stomata confined to the abaxial surface, crystals in the form of druse and/or prismatic shapes, trichomes of the stellate, peltate, or simple cylindrical type, sheathing and supporting elements associated with the venation, and pinnate primary venation accompanied by campto- dromous secondary venation. Interestingly, almost all variation observed in the leaf structure of the ten smaller styracaceous genera is encountered among species of the large genus Styrax. Leaf characters that are not present in Styrax but which occur elsewhere in the Styracaceae include the distinct petiolar brachy- sclereids in Huodendron, the sclerenchymatous supporting sheath cells in Pamphilia aurea Martius (/rwin et al. 20974 and 29208) that are per- pendicular to the long axis of the veins, and the unique features of crystal sand, actinodromous venation, and petiole vasculature in Parastyrax. Al- though many foliar characters appear to be extremely useful in the de- limitation of taxa, a larger sample size is needed to confirm these observa- tions. Some of the anatomical variation lends itself to possible ecological in- terpretation, although the absence of detailed habitat data for many of the specimens observed makes such interpretation difficult. In coriaceous leaves, such as those of Brazilian Pamphilia, there does appear to be a strong correlation between more intensive sclerification along the veins and veinlets, the presence of inrolled leaf margins, thicker cuticles on both eee compact spongy mesophyll, and an extremely dense indumentum the abaxial surface of the leaves. Many of these characters have been 1979] SCHADEL & DICKISON, LEAF ANATOMY 19 associated in other families with adaptations to a dry environment and may represent similar adaptation by styracaceous species. In leaves of species typically found in a mesic environment, such as Halesia (which occurs naturally in southeastern U. S. and China), there appears to be a strong correlation between less sclerification along the veins and veinlets, thin- ner cuticles on both surfaces, abundant air spaces in the spongy mesophyll, and less dense indumentum Relative levels of specialization of the foliar venation of woody dicots have been determined through comparative morphological studies of fos- sil and extant leaves (Hickey, 1971; Hickey & Doyle, 1972; Hickey, 1973; Hickey & Wolfe, 1975; Hickey & Doyle, 1977). Trends of specialization are represented by: 1) an increase in regularity of both low and high order venation; 2) a shift from indistinct vein orders to vein orders that are readily distinguishable from one another; 3) a shift from tertiary veins that follow a random course to tertiary veins that are very regular, often with rigid percurrent veins traversing the intercostal areas; and 4) a pro- gressive development from random, incomplete areolation to orthogonal, well-developed (perfect) areolation. The venation of the Styracaceae is predominantly at an advanced level of organization in accordance with the above trends. The secondary veins are generally regular in course, and the tertiary veins linking the secondaries demonstrate trends toward being nu- merous, well differentiated, percurrent, and regularly spaced in a close arrangement. Another index of advancement is the formation of well- developed areoles of relatively constant size and placement in which vein- lets are either absent or short and unbranched as in Huodendron, Para- styrax, Sinojackia, and several species of Styr ax. Petiole vasculature of the Styracaceae is quite diverse and, with the ex- ception of that found in Parastyrax, consists of an arc, an arc with in- vaginated ends, a medullated cylinder, or a medullated cylinder with medul- lary tissue; these characters may or may not be accompanied by cortical bundles. In the leaves of several species of Styrax, the occurrence of both medullary and cortical petiolar vasculature is correlated with compact spongy mesophyll and an increase in leaf coriaceousness. Whereas each of the styracaceous genera except Styrax is characterized by a single vascular configuration, all observed patterns of petiole vasculature occur among species of the large genus Séyrax. The presence in the petiole of Parastyrax of a highly dissected cylinder of collateral bundles that is accompanied by numerous medullary bundles and two smaller dorsal dissected cylinders is exceptional for the family. Minute, crenulate teeth are present in all genera investigated except Huodendron and Pamphilia. Each tooth is vascularized by a medial vein that is usually slightly expanded at the base and tapered near the apex. A glandlike cap composed of thin-walled cells occurs at the apex of each tooth. Hickey and Wolfe (1975) described what they termed the “Theoid Tooth and its inferred variants” as present in the Ebenales, but it is not certain whether the typical styracaceous tooth morphology observed in this study falls within the limits of variation observed in that survey. 20 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60 Although trichomes have traditionally been employed for systematic purposes in the Styracaceae (Perkins, 1907; Standley & Steyermark, 1940; Gonsoulin, 1974), it is clear that their morphological diversity has not yet been satisfactorily described. Among stellate trichomes, enormous varia- tion within the family can be observed in such features as size, color, dis- tribution, mode of attachment, appressed as opposed to stalked habit, number of arms, thickness of arms, degree of lateral fusion of arms, and number and orientation of cells composing each arm. Because of the large amount of gradation between these features, the value of stellate trichomes as diagnostic characters for separating taxa within the family is uncertain until these trichomes are more accurately characterized by employing a wider range of materials. The same difficulties are also evident, although to a much lesser extent, among peltate scales and simple cylindrical tri- Baas (1972) presented evidence for the removal of A frostyrax from: the Styracaceae and for its subsequent placement in the malvalean family Huaceae. The present study affirms these conclusions. The number of characters separating the Huaceae from the Styracaceae is substantial. Characters distinguishing A frostyrax leaves from styracaceous foliage in- clude paracytic stomata, extremely well-developed, rectangular areolae, the presence of caducous stipules, and fimbriate marginal ultimate vena- tion. According to Baas (1972), additional features distinguishing the Huaceae from the Styracaceae include the choripetalous corolla (vs. sym- petalous in the Styracaceae), the 3-porate pollen (vs. 3-colporate), the trilacunar nodes (vs. unilacunar), and the cork arising in subepidermis (vs. in pericycle). The genus Lissocarpa was removed from the Styraca- ceae by Perkins (1907) and has subsequently been regarded as constitut- ing the closely related monogeneric family Lissocarpaceae (sensu Engler & Gilg, 1924; Hutchinson, 1926, 1959, 1967, 1969, 1973). In addition to reproductive characters, features of foliar anatomy that can be used to separate Lissocarpa from the Styracaceae include diffuse vesiculose sclereids in the petiole and mesophyll, petiole vasculature consisting of an arc with outwardly curving ends, and fimbriate marginal ultimate venation. The genus Parastyrax has a number of typical styracaceous floral char- acteristics (bisexual flowers axillary on new shoots; calyx subentire; petals 5, imbricate; stamens 10; anthers introrse; ovary usually inferior) but also has some leaf characters that are markedly different from the other styracaceous genera. These unique leaf characters include a distinctive petiolar vasculature, actinodromous primary venation, and the presence of crystal sand in the leaf mesophyll. Additional points of difference be- tween Parastyrax and other taxa of the Styracaceae include its unusual height of 150 feet or more and its drupaceous fruit with a glabrous, fleshy exocarp marked with elongate, whitish lenticels (Smith, 1920). These dis- tinguishing characters indicate that further intensive investigation of Parastyrax is warranted to discern more accurately its affinities with other styracaceous genera. According to Wood and Channell (1960), the genus Halesia is most 1979] SCHADEL & DICKISON, LEAF ANATOMY 21 closely related to Pterostyrax, Rehderodendron, Sinojackia, and Mellio- dendron. These five genera share a number of similar reproductive char- acters (including ovary two-thirds to completely inferior, four to eight ovules in each loculus, a single elongate style, and indehiscent fruit), as well as a substantial number of common leaf characters. According to Gon- soulin (1974), the genera Alniphyllum and Pterostyrax are most closely related to Styrax. Alniphyllum differs from Styrax mainly in its winged seeds and filaments connate distally to form a tube, whereas Pterostyrax differs from that genus primarily in its two-thirds to completely inferior ovary. These three genera share such reproductive characters as bisexual flowers, campanulate calyx, five corolla segments or free petals, ten stamens, and linear anthers. While it is true that there is a high correlation between the leaf characters of Alniphyllum and Pterostyrax and those of several species of Styrax, it is also true that almost all of the variation observed in the leaf characters of the ten small genera occurs in the large genus Styrax. ACKNOWLEDGMENTS We are grateful to Dr. Albert E. Radford and Dr. Patricia G. Gensel for their attentive consultation during the course of this study and for their careful review of the manuscript. We would like to express our gratitude to the following persons and institutions for kindly providing leaf specimens: Dr. Richard H. Eyde and Mr. James J. White, III, Smithsonian Institution, Washington, D. Cs Dr. Stephen A. Spongberg, Arnold Arboretum, and Professor Norton G. Miller, Arnold Arboretum and Gray Herbarium, Harvard University, Cam- bridge, Massachusetts; Dr. Jimmy R. Massey, NCU Herbarium, Chapel Hill, North Carolina; Dr. Pieter Baas, Rijksherbarium, Leiden, Nether- lands; and especially Dr. D. M. Henderson, Royal Botanic Garden, Edin- burgh, Scotland, for sending material of the genus Parastyrax. LITERATURE CITED Arnott, H. J. 1959. Leaf clearings. Turtox News 37: 192-194. Baas, P. 1972, Anatomical contributions to plant anatomy II. The affinities of Hua Pierre and Afrostyrax Perkins et Gilg. Blumea 20: 161-192. BENTHAM, G., & J. D. Hooker. 1876. Gen. Pl. 2: 666-671. Bessey, C. E. 1915. The phylogenetic taxonomy of flowering plants. Ann. Missouri Bot. Gard. 2: 109-164 Cronouist, A. 1968. The evolution and classification of flowering plants. 396 pp. Houghton-Mifflin Company, New DAHLGREN, R. 1975. A system of classification of the pa aime is be used to demonstrate the distribution of characters. Bot. Not. 128: —147. EncLeR, A., & E. Gite. 1924. Syllabus Pflanzenfam. ed. 2. p. ae ae Borntraeger, Berlin. Faun, A. 1974. Plant anatomy. ed. 2. 611 pp. Pergamon Press, New Yo Gonsoutin, G. J. 1974. A revision of Styrax (Styracaceae) in North eee Central America, and the Caribbean. Sida 5: 191-258. a2 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60 GuRKE, M. 1891. Styracaceae. 7x: A. ENGLER & K. PRANTL, eds., Nat. Pflan- zenfam. IV. 1: 172-180 Hickey, L. J. 1971. Evolutionary aay of leaf a features in the woody dicots. Am. Jour. 58: 469. | Abstrac . 1973. Classification of the ue of ae hae leaves. /bid. 60: 17-33. & J. A. Doyle. 1972. Fossil ev sar on evolution of angiosperm leaf venation. Am. Jour. Bot. 59: 661. aa 1977, Early coe “fossil evidence for angiosperm evolution. Bot. Rev. 43: 3-104. & J. A. Wotre. 1975. The bases of angiosperm phylogeny: vegetative morphology. Ann. Missouri Bot. Gard. 62: 538-590. Hoitmcren, P. K., & W. KevuKEN. 1974. Index aaa Part 1. The herbaria of the world. ed. 6. Reg. Veg. 92: 1-39 Huser, H. 1963. Die Verwandtschaftsverhaltnisse a ‘Rosifloren. Mitt. Bot. Staatssam. Miinchen 5: 1-48 HUTCHINSON, J. 1926. The families of flowering plants. Vol. 1. 328 pp. Mac- millan and Co., Ltd., London. 1959. The families “ot flowering plants. ed. 2. Vol. 1. 510 pp. The Clarndon Press, Oxford. 1967. The genera of flowering plants. Vol. 2. 659 pp. The Claren- don Press, Oxford. 1969. Evolution and phylogeny of flowering ee Dicotyledons: facts and theory. 717 pp. Academic Press, New 1973. The families of flowering plants: ed. a 968 pp. The Clarendon Press, Oxfor JOHANSEN, D. A. 1940, Plant microtechnique. 523 pp. McGraw-Hill, New BS rk. LEE, S. 1973. Forest botany of China supplement. 477 pp. Chinese For- estry Association, an Means 5, A. 1972. Phylogenetic, biochemical, and morphological studies with some living specimens of naturally extinct or relictual Styrax species. Bar- tonia 41: = 9. = C. R., & L. CHatk. 1950. Anatomy of the dicotyledons. Vol. II. 500 pp. a Clarendon Press, Oxford. ae K 1936. Determination of the Formosan species of Styracaceae based on the morphological characters of leaves. Jour. Taihoku Soc. Agr. For- estry 1: 66-76. Nicotson, D. H., & G. C. STEYSKAL. 1976. The masculine Aes of the gen- eric name Styrax Linnaeus (Styracaceae). Taxon 25: 581-587 Nootespoom, H. P. 1975. Revision of the ee ee of the Old World, New Caledonia excepted. Leiden Bot. Ser. 32) PERKINS, J. 1907. Styracaceae. Ju: A. ENGLER, Pflanzenr. IV. 241: 1-111. 1909. Eine neue Gattung der Styracaceae aus dem tropischen Afrika. ae Bot. Jahrb. 43: 214-217. 928. Ubersicht iiber die Gattungen der Styracaceae sowie Zusammen- ee der Abbildungen und der Literatur iiber die Arten dieser Familie bis zum Jahre 1928. 36 pp. W. Engelmann, Leipzig Rao, T. A., & O. P. BuupaL. 1973. Typology of sclereids. Proc. Indian Acad. 5 ares REVEAL, J. L., & M. J. Setpin. 1970. On the identity of Halesia carolina L. (Styracaceae). Taxon 25: 123-140 1979] SCHADEL & DICKISON, LEAF ANATOMY 23 Sarcar, G., & M. L. oo 1975. Foliar anatomy of two species of Styrax. Geobins 2: 110-11 SMITH, W. W. 1920. ce on certain Asiatic Styracaceae. Notes Bot. Gard. Edinburgh 12: 231-236. SOLEREDER, H. 1908. Systematic anatomy of the dicotyledons. (English trans- lation by L. A. Boopte & F. E. Fritscu.) Vols. 1, 2. 1182 pp. Oxford Univ. Press, London SPONGBERG, S. A. 1976. "Styracaceae hardy in temperate North America. Jour. Arnold Arb. 57: 54-73. STANDLEY, P. C., & J. A. STEYERMARK. 1940. Styracaceae. /n: sey of Cen- tral American plants. I. Field Mus. Publ. Bot. Ser. 22: 221-32 eet C. G. G. J. van. 1932. Contributions a Vétude de la flore as Indes néerlandaises. KXI. The Styracaceae of Netherlands India. Bull. Bot. Gard. Buitenzorg III. 12: 272. TAKHTAJAN, A. L. 1969. Flowering plants: origin and dispersal. (Translated from Russian by C. JEFFREY.) 310 pp. Oliver & Boyd, Edinburgh . 1973. Evolution und Ausbreitung der Blutenpflanzen. 189 pp. Gustav Fischer Verlag, Stuttgart. THORNE, R. 1968. Synopsis of a putatively phylogenetic classification of the fowooer plants. Aliso 6: 57-66. 1976. A Se ae of the Angiospermae. /z: M. K. HECHT, W. C. STEERE, & B. WALLACE, eds., Evolutionary biology. Vol. 9. vii + 458 pp. Plenum Press, New Yor ee G. 1964. Styracaceae. In: A. ENGLER, Syllabus Pflanzenfam. ed. Band 2. pp. 400, 401. Gebriider Borntraever, Berlin. WILLIS, GeO 73: A dictionary the ia plants and ferns. ed. 8. Re- vised by H. K. Artry SHAW. 1245 + lxvi pp. Cambridge Univ. Press, Cambridge. Woop, C. E., & R. B. CHANNELL. 1960. The genera of the Ebenales in the southeastern United States. Jour. Arnold Arb. 41: 1-35 DEPARTMENT OF BOTANY THE UNIVERSITY OF NORTH CAROLINA CHAPEL HILi, NorTH CaROLINA 27514 APPENDIX. Specimens examined. Afrostyrax kamerunensis Perk. Cameroons: J. Mildbraed ee A*); G. Zenker 3922 (us). A erm Mildbraed. Congo: G. Gilber 10222 (a), J. Louis 3965 (us Alniphyllum eberhardtii Cuillaumir: Indochina: A. a 4490 (us); E. Poilane 19043 (A). A. fortune ae Chimen, China . Ching 3105 (A). A, hainanense Hay. Hainan ina: C. J. Let 442 fos terospermum Mat- sum. Formosa: £. H. W Deon 9992 (us); label sae illegible (us #1597269). Bruinsmia Le alee Boerl. & Koord. New Guinea: M. S. Clemens 2970 (a). North B > J. & M.S. Clemens 50339 be Halesia ere L. Jackson Co., North Carolin ne. Nesom s.n. (NCU). South Carolina: H. D. House 1866 (us). H. diptera var. diptera Ellis. Jefferson Co., * Herbarium abbreviations follow Holmgren and Keuken (1974). 24 JOURNAL OF THE ARNOLD ARBORETUM [voL. 60 ae oe W. B. ne ay (us). H. diptera ae - magniflora God- Tallahassee, Florida: R. AK. Godfrey 71367 (us). H. macgregori: Chun. Chekiang. China: Ching a a ) Huodendron biaristatum (W. W. aoe Rehder. Indochina: Petelot 6339 (us). H. chunianum Hu. Yunnan, China: H. T. Tsai 60571 (A). H. tibeticum (Anthony) Rehder. Tibet: J. PF. Rock 22020 (Us) Lissocarpa benthamii Gurke. British Guiana: H. A. Gleason 724 (GH). L. guianensis Gleason. British Guiana: J. S. de la Cruz 3076 (GH); N. Y. Sandwith 1590 (us Malad: endron wangianum Hu. Szechwan, China: Chiao & Fan 364 (a). M. xylocarpum Handel-Mazz. Kwangsi-Chuang, China: Ching 7132 (us). Kwang- tung, China: W. T. Tsang 26363 (a). Pamphilia aurea Martius. Brazil: H. S. Irwin, H. Maxwell, & D. C. Wass- aa 20974 us); Irwin et al. 29208 (GH). P. pedicellata var. ovalis Perk. Braz Mexia 5778 (us). P. styracifolia A. DC. Brazil: J. E. de Oliveira s.n. (us #7103054). Parastyrax lacei W. W. Smith. Burma: J. H. Lace s.n. (£). Pterostyrax cavaleriei Chun. Kweichow, China: Y. Tsiang 6434 (wus). corymbosus Sieb. & Zucc. Japan: VY. Tateishi 7010 (us). P. hispidus Sieb. . Zucc. Konigl. Botanischer Garten, Breslau, Poland: C. Baenitz sn. (us #1155541). Rehderodendron indochinense Li. Yunnan, China: A. WM. Feng 12307 (aA). R. kweichowense Hu. Kwangsi, China: S. K. Lau 28706 (a). R. ena Hu. Szechwan, China: W. P. Feng 2663 (a); T. C. Lee — Sinojackia henryi (Dimmer) Merr. Szechwan, China: A. ae 8865 (GH). S. rehderiana Hu. Without location: oo unknown ee (L). 5S. xylocarpa Hu. Kiangsu, China: W. Cheng 4494 (v Styrax agrestis G. Don. Solomon islands: L. J. Brass 3261 (a). S. ambiguus Seub, Brazil: £. Warming s.n. (Us #1110459). S. americanus var. americanus Lam. Florida: PF. Rugel s.n. (us #512620). S. americanus Lam. var. pulveru- lentus Spares, Perk. Escambia Co., Alabama: E. S. Ford 5242 (us). Hardin 3: V. L. Cory 52817 (us). S. argenteus var. argenteus Presl. Nayarit, Mexico: J. G. Ortega 60 (us). S. argenteus Pres] var. hintonii (Bullock) Gon- soulin. Michoacan, Mexico: Bro. G. Arséne 8472 (us). S. argenteus Presl var. ramirezi (Greenman) Gonsoulin. Chiapas, Mexico: C. SS Purpus 157 (us). S. benzoin oo Java, Indonesia: Blume s.m. (a). Trinidad, West Indies: V Broadway sm. (us #1194642). S. camporum Pohl. Brazil: G. Hatsch- bach & O. Guimaraes 21776 (us); L. O. Williams 6432 (GH). S. cespedesii agi Colombia: £. L. Little, Jr. 8820 (us). S. conterminus Smith. El Salvador: H, Allen 7184 (us). S. crotonoides Clarke. Singapore: Field 36937 (a). S. davillifolins Perk. Venezuela: A. Jahn s.n. (Us #1189096). S. faberi Perk. China: C. O, Levine 467 (GH). S. fanshawei Sandwith. British Guiana: D. B. Fanshawe 5607 (us). S. ferrugineus Nees & Martius. Brazil: Irwin et al. 13659 (us); G. O. A, Malme 1958a (cH). S. formosanus Matsumura. Formosa: Wilson 10266 (A). S. glaber Swartz. St. Kitts, West Indies: R. K. Wadsworth 495 (a). S. glabra- tus Schott. Brazil: A. Loefgren 4165 (us). S. glabrescens var. glabrescens Bentham. Mexico: Purpus 88 (us). S. glabrescens Bentham var. pilosus Perk. Mexico: £. Matuda 4179 (us). S. grandifolius awe Garland Co., Arkansas: C. F, Adams s.n. (us #£2420427). Gwinnett Co., Georgia: H. A. Allard 194 (vs). . oo Perk. Formosa: Wilson 9933 (a). S. een We Macbride. Juanjui, G. Klug 4282 (us). S. hypargyreus Perk. Venezuela: H. Pittier 15725 1979 | SCHADEL & DICKISON, LEAF ANATOMY 25 (US) 3: es ee Perk. Kwangsi, China: Tsang 24105 (A). S. Lae = Watson. Mexico: C. A. Pringle 11012 (us). S. japonicus Sieb. & Zucc. Jap N. Fukuoka 5852 (us); M. Hiroe 10251 (a). Washington, D. C.: G. B. ene sm. (us #478713). S. langkongensis W. W. Smith. Yunnan, China: Rock 10546 (A). S. latifolius Pohl. Brazil: ee unknown 7269 (us): H. M. Cur- ran 627 (cH). S. longifolius Standley. Peru: R. Ferreyra 4546 (us). S. macro- calyx Perk. Colombia: J. C. Mutis 2465 ao) S. macrotrichus Perk. Colombia: Mutis 5207 (us). S. martii Seub. Brazil: [rwin et al. 32264 (us). S. micrasterus Perk. Colombia: Mutis 4260 (us). S. microphyllus Perk. Colombia: Mutis 2558 (us). S. obtusifolius Griseb. Cuba: R. A. Howard 5618 (us); A. Gonzales 477 (A). S. ochraceus Urban. West Indies: E. L. Ekman 13883 (us); Bro. A. H. Liogier 11364 (cH). S. officinalis L. var. fulvescens (Eastw.) Munz & Johnston. Calli- fornia: L. P. Abrams 6273 (us); E. A. Purer 6680 (us). S. officinalis L. var. redivivus (Torrey) Howard. Placer Co., California: £. Crum 2043 (us) ovatus A. DC. Peru: Mexia 8171 (cH). iy pallidus A. DC. Venezuela: L. Wil- liams 10353 (A). S. paralleloneurus Perk. Sumatra, Indonesia: H. O. Forbes 2354 (GH). S. pearcei Perk. Bolivia: O. — 5953 (us). Styrax pentlandianus Remy. Bolivia: A. Miguel 2867 (us). se aL cei Perk. Chekiang ina: H. H. Hu 1689 (a). Kiangsu, China: i PF. Tsu s.n. (us #1346056). S ‘platani- folius Engelmann. Kendall Co., Texas: E. J. Parker 11474 (us). S. pohlii A. DC. Brazil: collector unknown 2885 (us). S. poissonianus Perk. Peru: C. Schunke A39 (us). S. portoricensis Krug & Urban. West Indies: collector unknown s.n. (us #404089). S. pseudargyrophyllus Sleumer. Colombia: L. Munoz s.n. (us #1950764). S. serrulatus Roxb. Tibet: G. H. Cave 1.9.1920 (a). S. subcordatus Moric. Brazil: Zrwin et al. 12490a (GH). S. subcrenatus Handel-Mazz. Hainan, China: F. C. How 71118 (a). S. suberifolius Hooker & Arnott. Kwangtung, China: Tsang 25927 (a). Hong Kong: W. Y. Chun 5071 (a). Taiwan: A. Hsieh n. (US #2063471). S. sumatranus J. J. Smith. Sumatra, Indonesia: Yates 2532 (a). S. yapobodensis (Idrobo & R. E. Schultes) Steyerm. Colombia: R. £. Schultes 14394 (A). EXPLANATION OF PLATES PLATE I Transverse sections of petiole and midrib eee patterns of vasculariza- tion. A, Afrostyrax kamerunensis (Zenker 3922): 1, medial petiole; 2, distal petiole; 3, midrib 2 mm. from petiole; 4, midrib 4 mm. from petiole; 5, medial midrib. B, Lissocarpa guianensis (de la Cruz 3076): 1, basal petiole; 2, medial petiole; 3, distal petiole; 4, medial midrib. Key to tissues: solid black, sclerenchy- ma; stipple, phloem; vertical lines, xylem, PLATE II Transverse sections of petiole and midrib illustrating patterns of vasculariza- tion. A, Parastyrax lacei (Lace s.n.): 1, medial petiole; 2, distal petiole; 3, midrib 5 mm. from petiole; 4, medial central actinodromous vein. B, Rehdero- dendron kweichowense (Lau 28706): 1, basal petiole; 2, medial petiole; 3, distal petiole; 4, medial midrib. Key to tissues: solid black, sclerenchyma; stipple, phloem; vertical lines, xylem. 26 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60 PLATE III Transverse sections of petiole and midrib illustrating patterns of vasculariza- tion. A, Pamphilia aurea (Irwin et al. 29208): 1, basal petiole; 2, medial petiole; 3, distal petiole; 4, midrib 2 mm. from petiole; 5, medial midrib. B, Simojackia xylocarpa (Cheng 4494): 1, basal petiole; 2, medial petiole; 3, distal petiole; 4, medial midrib. C, Styrax americanus (Rugel s.nm.): 1, basal petiole; 2, medial petio ale - distal petiole; 4, medial midrib. D, Styrax japonicus (Fukuoka 5852): 1, basal petiole; 2, medial petiole; 3, distal petiole; 4, medial midrib. Key to eee solid black, sclerenchyma; stipple, phloem; vertical lines, xylem. PLATE IV Transverse sections of petiole and midrib illustrating patterns of vasculariza- tion. A, Styrax benzoin (Broadway s.n.): 1, basal petiole; 2, medial petiole; 3, distal petiole; 4, medial midrib. B, Styrax micraster (Miieds 4260): 1, basal petiole; 2, medial oan 3, distal petiole: 4, medial midrib. C, Styvas ochra- ceus (Ekman 13883): 1, basal petiole; 2, me me pe a. 3, distal petiole; 4, medial midrib. Oe Sign pohliu (collector unknown 2885): , basal petiole; 2, medial petiole; 3, distal petiole; 4, medial midrib. Key i tissues: solid black, aa stipple, phloem; vertical lines, xylem. PLATE V Venation patterns (leaves not drawn to scale). A, Afrostyrax kamerunensis (Zenker 3922). B, Lissocarpa benthamii (Gleason 724). C, Alniphyllum eber- hardtu (Petelot 4490). D, Alniphyllum pterospermum (W oe 9992). E, Bruin- sma styracoides (J. & M.S. Clemens 50339). F, Halesia carolina (Nesom s.n.). G, Halesia macgregorii (Ching 2132). H, Huodendron tibeticum (Rock 2202 0). I, Melliodendron xylocarpum (Ching 7132). PLATE VI Venation patterns (leaves not drawn to scale). A, Pamphilia aurea (Irwin et al. 20974). B, Parastyrax lacei (Lace s.n.). C, Pterostyrax hispidus (Baenitz M. Rehderodendron indochinense (K. M. Feng 12307). E, Sinojackia asilocarie (Cheng 4494). F, Styrax agrestis (Brass 3261). G, Styrax argenteus var, argenteus (Ortega 60). H, Styrax faberi (Levine 467). 1, Styrax officinalis var, redivivus (Crum 2043). PLATE VII Leaf areolation. A, Afrostyrax kamerunensis (J. Mildbraed 10688), > 107, well-developed areoles. B, Huodendron biaristatum (Petelot 6339), * 40, ob- serve relatively consistent size and shape of areoles. C, Simojackia xylocarpa (Cheng 4494), X 27, relatively well-developed areoles. D, Styrax glabratus (Loefgren 4165), * 36, imperfectly developed areoles. E, Styrax japonicus (Fukuoka 5852), X 27, incompletely developed areoles. F, Styrax suberifolius (Chun 5071), X 27, observe branching of ultimate veinlets and imperfectly de- veloped areoles 1979] SCHADEL & DICKISON, LEAF ANATOMY 27 PLATE VIII Tooth morphology. A, B, Styrax serrulatus (Cave 1.9.1920): A, X 40, note vascularization of tooth by tertiary vein. B, X 107, observe expansion of vascu- lar tissue at base of tooth. C, Styrax japonicus (Fukuoka 5852), X 40, gland- like cap on apex of tooth. D, Styrax faberi (Levine 467), X 107, note tapering of vascular tissue at apex of tooth below glandlike cap. PLATE IX Transverse sections of lamina. A, Afrostyrax lepidophyllus (Gilbert 10222), note prominent papillate abaxial epidermis. B, Bruinsmia styracoides (hae. 2970), X 160, observe isodiametric shape of spongy parenchyma cells. C, Huodendron Tie es (Tsai 60571), X 160, prominent cuticular striations on both surfaces. D, Pamphilia aurea (Irwin et al. 29208), X 107, note upright cells in adaxial sheath extension. E, Styrax ferrugineus (Irwin et al. 13659), X 160, observe prominent adaxial bundle sheath extension. F, Styrax glaber (Wadsworth 495), X 160, note absence of bundle sheath Ee rencion: PLATE X Transverse sections of lamina. A, Styrax macrotrichus (Mutis 5207), * 265 observe weakly differentiated mesophyll. B, Styrax micrasterus (Mutis 4260), X 160, note compact spongy parenchyma. C, Styrax ochraceus (Ekman 13883), X 135, observe three layers of palisade parenchyma. D, Styrax philadelphoides (Tsu s.n.), - 160, bundle sheath extensions occurring in both adaxial and abaxial positions. Styrax portoricensis (Us #404089), X 265, observe weakly seme, mesophyll. F, Styrax pseudargyrophyllus (Murioz SN Se OO, note biseriate epidermis (hypodermis?). Jour. ARNOLD Ars. VOL. 60 UUiil 6 AN | | DY f/f | \ X Ny \ ‘ SUT amy Tl N LETT NA My cv Lif /\{ / + 2 Gad if | \ I} | NI Nii A mul PLATE I (qa) -) Nf || NU UW UVAVTRGAAIAIAI TT ) | VY ti | titi 1 600 4m. B SCHADEL & DickIson, LEAF ANATOMY PuatTeE II Jour. ARNOLD Ars. VOL. 60 2, 99 8000%o $8, 0% 3 a, NS . g® © 6) e © “09% ye | {l oa Ee QU Op : t eG e.2¢ EY g ‘ O82 @& 2A “2 SED Gl © =) / am LD cm Mh \ ‘e 9? \ ( 1a 7 Bi “6% U7 as ee m t S ue Pas e) @ 09 7 AN Uy 2 y ‘ap y (lh ES © @ @,,= | < ‘fa Con TI I ay) | 7 “Seq U pe Bo .__300um. A 1000um. 2 —s 7 SCHADEL & DIcKIsON, LEAF ANATOMY PuaTE III Jour. ARNOLD ARB. VOL. 60 = | + | —— za NX ( || \ Ne Pal y ThtT are | ASTTTTTTITTTTTT 7 SLU LL “ aD is (| lg milly VAT I [Jed \ Ui Mere Wy / SE MUTE | ee 2 (ly Alllil | t CAA 1 TW (7 J TN, | by | | = | \ f j 1T i | | ammmamn tll QO WE) ppp | | | La NO | |v a ALE |} | bya =~ 4 \ ] rr | \ | ®) (D oN | LLL’ atl Panel ~~ ee re ota ae AY} Do TD Vt | || | Sak LULL l? VW N\\y B ' 2 (\! | S any WN ETT 4 a wo ~ | Ley ae a re Wiisnnsnnn tania / LUTE EL Ee SCHADEL & DickIson, LEAF ANATOMY Jour. ARNOLD ARB. VOL. 60 PLATE IV aN Ly TT » a (( - 1] af | | y) Sere MULL [Less “uy t 2 ® @ iN in Tm mr AW : a A va WN mL | IO! | | iY /| AD) C3) \ | 1 \} UY L ) 1 Jl ‘alll for HWTTHITAAS mie i sa 2 SU wy = ie 2 @® t & 9 © me SCHADEL & DICKISON, LEAF ANATOMY PLATE V Jour. ARNOLD Ars. VOL. 60 SCHADEL & DICKISON, LEAF ANATOMY PraTE VI Jour. ARNOLD ARB. VOL. 60 SEE SP a, eS ae Sep Pe , y Ne i og moment WOT Ww? NA) we ; AT I EERE SO NTO RRR é EIDE RET S EU S) RISIs s EIEN eae ne oe = SQL jaen \ aI NSRINITEAN uw SOG S: SEI EES” < 4 LA RARER IR STS ROR Serine rerecs Wo Shs cX THEY ie SHY SL Ke ia Vai Perils STS SCHADEL & DickIson, LEAF ANATOMY Jour. ARNOLD Ars. VOL. 60 Puat_e VII SCHADEL & DICKISON, LEAF ANATOMY Jour. ARNOLD ARB. VOL. 60 Puate VIII SCHADEL & DICKISON, LEAF ANATOMY Jour. ARNOLD ARB. VOL. 60 PLATE IX re Ie, fo Ee EN EES MS : pled SS el eT gt est Ms ried» i ‘4 a3 bp ee one a § fe & ; tht 8 e *: 7 Cc D SCHADEL & DickIson, LEAF ANATOMY Jour. ARNOLD Ars. VOL. 60 PLATE X san eres SCHADEL & DICKISON, LEAF ANATOMY 38 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 A REEXAMINATION OF PORTLANDIA (RUBIACEAE) AND ASSOCIATED TAXA ! ANNETTE AIELLO THE OBJECTIVES of the present study have been to establish the generic limits of the genus Portlandia, to determine the position of taxa not prop- erly assigned to it, to determine the number of species belonging to Port- landia and to describe them, and, finally, to reexamine all of the important taxa ever placed in the genus for additional characters that may have been overlooked or underrated by past workers. DESCRIPTION OF THE FAMILY The Rubiaceae, containing approximately 6000 species and over 600 genera, is the third largest family of dicotyledons. It is a mostly tropical group of trees, shrubs, climbers, and herbs, which with very few exceptions have opposite, stipulate leaves and sympetalous, epigynous flowers. The inflorescence is fundamentally a cyme and varies from a much-branched cymose panicle to an inflorescence reduced to a solitary flower. The corol- la aestivation can be valvate, contorted, or imbricate. The fruit may be a capsule, a berry, a pyrene, or a schizocarp, and varies in number of locules and number of ovules per locule. Nectar is usually secreted by a gland around the base of the style. While heterostyly is common within the family, it is not found in all tribes. Cinchona, the source of quinine, and Coffea, the seeds of which are used to make coffee, are genera of eco- nomic importance. /xora, an evergreen shrub or small tree with red, orange, yellow, or white flower clusters, and Gardenia, with scented white flowers, are frequently cultivated in warm climates as ornamentals. Ac- cording to information gathered from Willis (1973), about 37 percent of the genera are monotypic. This high incidence may be due to a lack of understanding of the essential generic characters within the family; more likely, it may indicate that rapid evolution took place in the family at some time in the past and was followed by widespread extinction, leaving many genera isolated from their near relatives and resulting in a group that is, at the same time, clearly natural and extremely diverse. A major problem within the family has been that it contains a great many genera to belong to different tribes. * Extracted from a thesis submitted by Annette Aiello to the Department of Biology, Harvard University, in partial fulfillment of the requirements for the degree of Doc- tor of Philosophy. 1979] AIELLO, PORTLANDIA 39 HISTORY OF CLASSIFICATION WITHIN THE RUBIACEAE In 1763 Adanson divided the Aparines, as he called the group of taxa now known as the Rubiaceae, into two sections based on the number of ovules in their fruits. Section I, with two ovules, included seventeen genera, and section II, with more than two ovules, included fourteen. With a total of only thirty-one genera known, and with those so evenly divided as to the number of ovules, his classification seems an almost natural one based on information available at the time. Twenty-six years later, Jussieu (1789) used the name Rubiaceae when he published a classification more elaborate than that of Adanson. In it he recognized four main groups of genera instead of two: 1) fruit didy- mous (based on Rubia), 2) fruit with two ovules (based on Coffea), 3) fruit with many ovules (based on Cinchona), and 4) fruit with many locules (based on Guettarda). In addition, Jussieu recognized subdivisions of these taxa based on the number of stamens (4, 5, 6, or more), on whether the leaves were whorled or opposite, and on whether the flowers were aggregated or not. Although the systems of Adanson and Jussieu were simple and easy to use, they cut through now-accepted tribes and even genera and are thus no longer usable. Since the time of Adanson and Jussieu, new genera have been added to the Rubiaceae at the rate of almost 2.5 per year, yet until recently (Verd- court, 1958; Bremekamp, 1966) no important advances toward a really natural classification of the family had been made. Adanson’s system of sections (equivalent in rank to subfamilies) has been followed almost ee in major treatments of the family (de Candolle, 1830; Rich- , 1830; Endlicher, 1838; Schumann, 1891). De Candolle (1830) and Richard (1830) based tribes on the degree of dryness or fleshiness of the fruits, a division that also tends to cut through natural groups. Bentham and Hooker’s (1873) proposed division into three series represented a compromise between the systems of Adanson and Jussieu. The first step toward a more natural classification was taken by Verd- court (1958), who proposed the division of the family into three subfam- ilies based on a number of correlated characters, especially presence or absence of raphides, multicellular uniseriate vs. unicellular nature of the hairs, and presence or absence of albumin in the seeds. In addition, he pean shape of the stigma, number of ovules per locule, and testa sculpturing. Although Bremekamp (1966) divided the family into eight subfamilies, his classification and that of Verdcourt are in basic agreement on most major issues. They both recognize three main subfamilies (Rubioideae, Cinchonoideae, and Guettardoideae), but Bremekamp also splits off five smaller subfamilies based on the presence of such unique characters as i styloides (columnar crystals in cells). While these systems represent a large advance toward a more natural classification within the family, it 40 JOURNAL OF THE ARNOLD ARBORETUM [ VoL. 60 will doubtless be a long time before basic relationships are well under- stood. In general, recent workers distinguish tribes by such characters as habit, presence or absence of spines, stipule shape and size, perfect or dioecious flowers, inflorescence type, corolla aestivation, corolla symmetry, stamen insertion, stigma shape, number of ovules per locule, ovule attachment, dehiscence or indehiscence of fruit, degree of dryness or fleshiness of fruit, and whether seeds are alate or exalate. Much work remains to be done at the tribal level in the areas of palynology, wood anatomy, and fruit and seed morphology, where further information may yield characters of taxonomic value. POSITION OF THE CONDAMINEEAE The tribe Cinchoneae was designated by Richard (1830) as including plants with dry, 2-locular, many-seeded fruits, opposite leaves, and inter- petiolar stipules. As redefined by later workers, the Cinchoneae retained only those taxa with seeds both winged and vertical or ascending. In this tribe all three types of corolla aestivation (valvate, contorted, and im- bricate) occur. Two tribes that resulted from splitting the Cinchoneae were the Rondeletieae and the Condamineeae (which contains the genus Portlandia). In both of these tribes the seeds are exalate and horizontal. Corolla aestivation is valvate in the Condamineeae, and contorted or im- bricate in the Rondeletieae. While at first glance these three tribes appear well separated, pollen and wood studies indicate that this may not be the case. Verdcourt (1958, 1976) has simplified matters to some degree by including the Condamineeae in the Rondeletieae. The result, however, is a second tribe containing all three aestivation types. A clearer under- standing of these three tribes could undoubtedly be obtained if they were simultaneously revised after careful study of fruit, seed, pollen, and wood characters. Within the Condamineeae a variety of characters have been used to separate genera. These include shape, size, and persistence of stipules and calyx lobes, position of stipules, number of corolla lobes, shape of corolla tube in cross section, position of stamen insertion, type of anther de- hiscence, and type of capsule dehiscence. HISTORY OF THE GENUS PORTLANDIA The genus Portlandia was first described by Patrick Browne (1756) in his Civil and Natural History of Jamaica. There he used a polynomial to describe “the large-leaf’d Portlandia,’ but did not assign a Latin epithet. It is clear from his description and untitled illustration (¢. xi) that the plant he had in mind was Portlandia grandiflora, later briefly described, named, and credited to him by Linnaeus (1759), who also cited ¢. 77, in the tenth edition of Systema Naturae. Since neither worker mentioned a specimen, and because Linnaeus cited ¢. 77, that illustration must be the holotype although it was mistakenly engraved with six stamens instead 1979 | AIELLO, PORTLANDIA 41 of five. This error was corrected in the 1789 edition of Browne’s book, where the engraving was labeled Portlandia grandiflora. Portlandia was named by Patrick Browne in honor of Margaret, Duch- ess of Portland and patroness of natural history. The name Portlandia was also applied in her honor to a genus of clams (Mollusca: Pelecypoda) by Morch (1857). Portlandia grandiflora, with its large, showy flowers, was well known to early workers. Aiton (1789) included it in his Hortus Kewensis (1: 228) and noted that it was introduced into cultivation in England in 1775 by a Mr. Ellis. The plant was illustrated in Curtis’s Botanical Magazine (t. 286. 1795) and in Jacquin’s (1763) Selectarum Stir pium Americanarum Historia (t. 44). It is represented in the Ware Collection of Glass Flowers at Harvard University by glass model no. 565, created by L. and R. Blaschka (1895) in Hosterwitz, Germany. Portlandia was among the very earliest genera included in the Rubia- ceae and was one of 31 genera listed by Adanson (1763) in his influential treatment of the family. Being both an early genus and one with showy flowers, Portlandia almost naturally became a catchall for other rubia- ceous plants with large flowers. Superficial characters had a great deal of influence on how plants were classified, and it was only gradually that more basic characters such as fruit type and seed arrangement were con- sidered important. Portlandia became a temporary resting place for quite a number of plants. By 1889 it included ten species, only three of which truly belonged there. In 1918 the genus reached its largest size — 21 spe- cies. Taxa have been removed periodically as old generic limits have been more clearly defined and new genera recognized. At one time or another Portlandia has included various taxa now assigned to Coutarea, Bikkia, Exostema, Siemensia, Coutaportla, Schmidtottia, and Hintonia. GENERA PREVIOUSLY SEPARATED FROM PorTLANDIA. Plants having axil- lary flowers, straight corolla tubes, unflattened capsules, and wingless seeds were recognized by Aublet (1775) as Portlandia. Plants having ter- minal flowers, gibbous corolla tubes, strongly flattened capsules, and winged seeds were considered distinct from Portlandia, and Aublet based the tropical American genus Coutarea on them. Unfortunately, he applied a new epithet (speciosa) instead of the previously used Hexandra, and not until 1889 was the taxon correctly named Coutarea hexandra (Jacq.) Schumann. At present, Coutarea is in need of revision, and the number of species in the genus is uncertain. In 1923 Urban named three new genera, Siemensia, Coutaportia, and Schmidtottia, based on plants originally described as Portlandia. He dis- tinguished the monotypic Siemensia (endemic to Cuba) on the basis of leaf anatomy. Its leaves have resin-producing secretory cells as well as raphides and lack crystal sand (calcium oxalate) and the typical palisade and spongy mesophyll layers. In contrast, Portlandia leaves lack secretory cells and raphides but have both crystal sand and the typical palisade and spongy mesophyll layers. Urban separated Coutaportla (also monotypic; 42 JOURNAL OF THE ARNOLD ARBORETUM [voL. 60 endemic to Mexico) from Portlandia based on a number of characters: flowers 4-merous vs. 5- to 7-merous; calyx tube compressed vs. terete; capsule septicidal vs. loculicidal; placentae subquadrate and attached only to middle of septum vs. linear and attached to septum for its whole length; ovules few and not completely covering placentae vs. many and cover- ing placentae; seeds unappendaged vs. appendaged (funicle persistent) ; and stipule margins denticulate vs. entire. Schmidtottia (ten species en- demic to Cuba) was distinguished as having terminal inflorescences, septi- cidal capsules, oval placentae that are attached only to the middle of the septum, unappendaged seeds, and stipules that are truncate and connate into a sheath (as opposed to triangular and distinct). PROPOSED CLASSIFICATION OF PoRTLANDIA. The genus Portlandia (Rubia- ceae) as redefined in this study is endemic to the island of Jamaica and comprises five species. Of the more than thirty taxa formerly included in Portlandia, sixteen still remained at the outset of the present study. Of these sixteen, only five are here accepted as Portlandia, four represent the three new genera Cigarrilla, Cubanola, and Thogsennia, one is assigned to Coutarea, three belong in /sidorea, two are conspecific with P. platantha, and one remains unassigned to a genus until the fruits are known. In addition, two other new genera, Ceuthocarpus and Osa, are separated from Schmidtottia and Hintonia, respectively. For a complete list of taxa as- sociated with Portlandia, see TABLE 1 TABLE |. List of taxa associated with Portlandia. Ceuthocarpus Aiel C. involucratus ( oe ernham) Aiello Cigarrilla Aiell C. mexicana on & Martius ex DC.) Aiello. Coutaportla Urban, Symb. Antill. 9: 146. 1923. C. ghiesbreghtiana (Baillon) Urban, ibid. 147. Coutarea Aublet, Pl. Guian. 1: 314. 1775 C. hexandra (Jacquin) Schumann in Martius, Fl. Brasil. 6(6): 196, 1889, Cubanola Aiello. C. daphnoides (R. Graham) Aiello C. domingensis (Britton) Aiello. Hintonia Bullock in Hooker’s Ic. Pl. 33: 1. 1935. atiflora (Sessé & Moc. ex DC.) Bullock, ibid. 4. H. lumaeana (Baillon) Bullock, ibid. 5. Tsidorea A. Richard ex DC. Prodr. 4: 405. 1830. I. brachycarpa (Urban) Aiello 1. oblanceolata (Urban) Aiello. I. polyneura (Urban) Aiello. Schmidtottia Urban, Symb. Antill. 9: 137. 1923. S. elliptica (Britton) Urban, ibid. 139, 1979] AIELLO, PORTLANDIA 43 _nitens (Britton) Urban, zbid. 141. », sessilifolia (Britton) Urban, ibid. 142. ». sessilifolia subsp. shafert (Standl ey) Borhidi, Bot. Kozlem. 58: 176. 1971. ’. uliginosa (Wernham) Urban, Symb. Antill. 9: 139. 1923 HAHN Siemensia Urban, Symb. Antill. 9: 143. 1923. S. pendula een Urban, ibid. 145. Thogsennia Aie ES eee a Richard) Aiello. MORPHOLOGY LEAVES Many of the leaf characters discussed below are summarized in TABLE The leaves of the 12 genera studied are all typical of the Rubiaceae in that they are opposite, simple, entire, and stipulate. Exceptions are Ceuthocarpus involucratus and Schmidtottia cubensis, with leaves arranged two or three per node. Six genera (Ceuthocarpus, Cigarrilla, Coutaportla, Isidorea, Portlandia, and Schmidtottia) have coriaceous leaves; those of some species of /szdorea are rigid and pungent as well. The leaves of Coutarea, Hintonia, Cubanola, Osa, Siemensia, and Thogsennia are chartaceous. When dry, the leaves of Cigarrilla and Thogsennia curl longitudinally; those of Ceuthocarpus and Schmidtottia are very dark (dark brown to black) above and much paler (green to pale brown) below. The hairs on the leaves of Ceuthocarpus and Schmidtottia are simple (FicuURE 1), while those on the leaves of Siemensia (FIGURE 2) are uni- seriate and have two to four cells each. In both types the hairs are usu- ally surrounded at the base by a rosette of epidermal cells. The remaining nine genera have glabrous leaves, although in Hintonia standleyana the leaves are pubescent when young. The venation in all cases is brochidodromous (as defined by Hickey, 1973) or co-arcuate (as defined by Melville, 1976), depending upon whose terminology is used. The secondary veins are alternate, with four to seven pairs per leaf, except in /sidorea where they range from ten to thirty in number and are closely packed together. The venation of /szdorea approaches Melville’s (1976) definition of “paxillate,” but the arches formed by successive secondary veins are still evident. In a number of species of Jsidorea, the very numerous higher order veins are not re- stricted to a single plane and almost entirely fill the thick leaves. Several species of Portlandia and Schmidtottia, as well as one of /sidorea, have sessile leaves; the rest have petioles of varying length. The largest leaves are found in Cubanola, Osa, Portlandia, and Thog- sennia; those of the other genera are intermediate to small in size. Leaf shape is quite variable, ranging from linear-lanceolate through el- liptic, ovate, and obovate, to nearly circular. The apices vary from long- TABLE 2. Summary of leaf shape and size in Portlandia and associated genera. GENUS PETIOLE LENGTH (mm. ) BLADE SHAPE BLADE LENGTH (cm. ) Ceuthocar pus Cigarrilla Coutaportla Coutarea Cubanola Hintonia Tsidorea Osa Portlandia Schmidtottia Siemensia Thogsennia ra 5-13 narrowly elliptic to elliptic-obovate or narrowly obovate oblong elliptic ovate to elliptic ovate, elliptic- oblong, or elliptic circular, elliptic, oblong-lanceolate, or obovate elliptic or oblong-elliptic elliptic-lanceolate or oblong-lanceolate 2-6 BLADE WIDTH (cm. 0.62 55 0.2-6.5 105 4.5-6.5 BLADE APEX BLADE BASE rounded acute, mucronate acute acuminate acute to acuminate acute to acuminate acute, obtus rounded, or erally acuminate; pungent long-attenuate rounded, obtuse, acute, or acuminate rounded, obtuse, acute, or gradually acuminate acute to acuminate acute to acuminate acute to attenuate obtuse to acute acuminate acute to obtuse acuminate to acute, or obtuse acute rounded, obtuse, narrowing to base, acute, or acuminate long-attenuate attenuate, Se acute, obtuse, or cordate rounded, obtuse, acute, nar rowed, subcordate. or cordate acute to obtuse acute bP WOLAYOUUV GIONUV AHL AO IVNAAOL 09 “10A] 1979] AIELLO, PORTLANDIA 45 attenuate through acute and obtuse to rounded, and the bases from at- tenuate through acute and obtuse to cordate. Epidermal cell anticlinal walls are undulate in Hintonia, Isidorea, Port- landia, and Osa, and slightly undulate to straight in the remaining genera. f (1 ae sue FIGURES 1, 2. hairs: 1, Schmidtottia elliptica (Leén 21131 (Ny)); 2, Siemensia pendula (Shafer 13411 (Ny)). Bar scales = 0.1 mm The stomata in all cases are paracytic, although the number of subsidiary cells varies from two in Cigarrilla, Coutaportla, Coutarea, Hintonia, and Portlandia, and two to three in Cubanola, Isidorea, and Thogsennia, to two to four in Siemensia, and four in Ceuthocarpus, Osa, and Schmidtottia. In Siemensia the guard cells are often sunken and difficult to see. Druse crystals were noted in cleared leaves of Cigarrilla, Hintonia, [si- dorea, Portlandia, Siemensia, and Thogsennia. Bundles of raphide crystals were found in the leaves of Siemensia. Leaves of Osa were not available for clearing. A red substance occurs in leaf cells of Ceuthocarpus, Coutarea, Schmidt- ottia, and Siemensia; however, the exact location of those cells was not determined. A study of leaf anatomy for the 12 genera covered here, and a general survey of leaf anatomy for the family Rubiaceae, would doubtless be a most profitable undertaking and might help to clarify tribal and generic relationships. Indeed, Urban (1923), when describing the genus Siemensia as distinct from Portlandia, did so on the basis of leaf anatomy. v 46 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 60 NopeEs Sinnott (1914) stated that, “The Rubiaceae are overwhelmingly uni- lacunar, the only exception observed being the genus Sarcocephalus. . . .” Howard (1970), after his survey of nearly ninety genera (pers. comm.) in the Rubiaceae, reported trilacunar nodes in twelve genera (including Port- landia) representing eight tribes of that family. Further work in the Ru- biaceae may reveal that the unilacunar node is the predominant type in the subfamily Rubioideae (sensu Bremekamp, 1966), with both nodal types being common in the Ixoroideae and Cinchonoideae (Howard, un- published data). Material of Coutaportla, Cigarrilla, and Portlandia (seedlings) was ex- amined for nodal type. In Coutaportia the nodes are unilacunar, with the single crescent-shaped trace giving off stipule traces just before it enters the petiole. From there it continues as a single trace to the base of the blade. Cigarrilla has trilacunar nodes, with the lateral traces leaving the stele at a point almost 10 mm. below the attachment of the leaves, while the median trace does not leave until immediately before it enters the petiole. It is at this later point that the lateral traces converge into the ends of the median trace, with portions going to the stipules. Within the petiole base the traces separate again and ered an full length of the petiole as a crescent-shaped trace plus two rib t Portlandia also has trilacunar ae (Howard, 1970), but differs from Cigarrilla in that the three traces leave the stele at approximately the same level. Examination of Portlandia grandiflora seedlings 1 ene that each of the opposite cotyledons is supplied with two traces from one gap, a pat- tern previously reported (Thomas, 1907) for seedlings ce many dicotyl- edons STIPULES Among the Sympetalae only members of the Rubiaceae have stipules as a constant character. These may be interpetiolar, intrapetiolar, or sheath- ing. By far the most common type for the family is the interpetiolar stipule. Often within this type, the adjacent members of the two pairs are united along their outer margins, resulting in two interpetiolar stipules per node. Among the twelve genera studied, Schmidtottia has truncate sheathing stipules (FicURE 3), Ceuthocarpus has intrapetiolar erose stipules (F1c- URE 4); the other ten genera have triangular ones. In all genera except Coutaportla, in which the stipules are free from each other, the two stip- ules at each node are at least slightly connate at their bases and thus en- circle the stem. In Coutaportla (FicuRE 5) and Hintonia (Ficure 6) the stipules are acute, while in Portlandia (FicurE 7) they are acute or obtuse. Cigarrilla 1979 | AIELLO, PORTLANDIA 47 13 14 Figures 3-14. Stipules: 3, Schmidtottia nitens; 4, Ceuthocarpus involucra- tus; 5, Coutaportla ghiesbreghtiana; 6, Hintonia latiflora; 7, Portlandia micro- sepala; 8, Cigarrilla mexicana; 9, Coutarea hexandra; 10, Cubanola domingen- sis; 11, Osa pulchra; 12, Siemensia pendula; 13, Thogsennia lindeniana; 14, Isidorea leptantha: A, before splitting; B, after splitting. Bar scale = 2 mm. 48 JOURNAL OF THE ARNOLD ARBORETUM [voL. 60 (FicurE 8), Coutarea (FicuRE 9), Cubanola (F1GuRE 10), Osa (FIGURE 1), Siemensia (FicuRE 12), and Thogsennia (Ficure 13) have acuminate to attenuate stipules. /sidorea (F1curE 14) differs from all the rest in hav- ing narrow, aristate stipules that eventually split from the base to the apex, giving the appearance of four stipules per node. At the same time, a groove opens in the stem, beginning just below each stipule and con- tinuing down to the next node. This arrangement gives the branches a distinctive appearance. The stipules of Cigarrilla and Coutarea also oc- casionally split from the base to the apex with age, but these two genera lack the stem groove of /sidorea. INFLORESCENCE Within the Rubiaceae the inflorescence is typically cymose, with the most common condition being a much-branched cymose panicle. Solitary terminal flowers are rare. In Ceuthocarpus, Coutaportla, Schmidtottia, and Siemensia the inflo- rescences are terminal. Ceuthocarpus is unique among these in having soli- tary terminal flowers, each surrounded at the base by bracts and perhaps representing the terminal flower of a reduced cyme. In Coutaportla and Schmidtottia the flowers are in terminal simple or reduced cymes. In sev- eral species of Schmidtottia these cymes are condensed, with the sessile flowers crowded together. Siemensia inflorescences are terminal on axillary shoots. Typically, there are seven flowers, of which four arise from the axils of the upper two pairs of leaves, and three form a simple terminal cyme. After flowering, the leaves fall off the entire shoot, leaving a long- stalked panicle with seven stalked capsules. Cigarrilla, Cubanola, Hintonia, Isidorea, Osa, and Thogsennia have the flowers arranged one (rarely three) per axil in the upper one to four pairs of axils. /sidorea differs from the other genera in that three species (J. brachyantha, I. leonardii, and J. leptantha) have the flowers borne on axillary short shoots. Short shoots appear on occasional specimens of other Isidorea species as well. This genus needs field study with respect to its growth habits. Portlandia and Coutarea have their flowers arranged in reduced cymes of one to six and one to three flowers per axil, respectively, in the upper one to four pairs of axils. In Portlandia there is variation in the degree of cyme reduction among the species. Portlandia harrisii has three to six flowers per axil in condensed cymes with pedicels 1-5 mm. long. Portlandia coccinea and P. platantha have reduced cymes with one to three (most commonly two) flowers per axil. Portlandia grandiflora has reduced cymes with one or two (most commonly one) flower(s) per axil. Often these single flowers bear an aborted bud near the base of the pedicel. Portlandia microsepala has one flower per axil. FLowers. While heterostyly is quite common in the Rubiaceae, it has not been reported for any genera within the Condamineeae and was not 1979] AIELLO, PORTLANDIA 49 a C — 17 YW \ 16 Ficures 15-19. Flowers, stamens, and styles: 15, Portlandia platantha; 16, P. grandiflora (Adams 13213 (uewn): ee ae microsepala: 18, P. coccinea (Aiello 1308 (A)); 19, P. harrisit (Aiello 1306 (A)). Bar scales = 10 mm. 50 JOURNAL OF THE ARNOLD ARBORETUM [voL. 60 found in any of the taxa included in this study. In all 12 genera examined, the stamens are attached to the basal 1 to 3 mm. of the corolla, and at anthesis both the style and the stamens reach to the sinus of the corolla lobes (except in Hintonia, Coutarea, and Ceuthocarpus, where they are exserted ). Few or no data on fragrance are available for Ceuthocarpus, Isidorea, Osa, Schmidtottia, or Thogsennia. Flower fragrance, where known, is as- sociated with wht or pink flowers. Calyx, corolla, stamens, and style are discussed separately in the sections that follow. Flowers of all genera studied are pictured in Frcures 15-31. Catyx. In all 12 genera studied, the calyx lobes or sepals are joined only at the very base, and in all except Coutarea and Coutaportla (which have deciduous sepals) they are persistent or marcescent on the fruit. The sepals of Osa, Thogsennia, and Cubanola are long (2—2.5 cm.) and subulate; those of Hintonia and Coutarea are shorter (0.7—1.5 cm.) but also subulate. Cubanola has keeled sepals, with the keels continuous with the wings of the capsule. The sepals of Coutarea are pubescent on their inner surface. The sepals of Ceuthocarpus, Siemensia, and Schmidtottia are linear to subulate, while those of /sidorea are stiff, needle shaped, and keeled, each with a very sharp tip. In Cigarrilla and Coutaportia the sepals are narrowly triangular. Each sepal of Coutaportla bears two or three red-tipped glands along each side near the base (FIGURE 32). With the exception of Portlandia microsepala, which has subulate sepals 2-9 mm. long and 0.5 mm. wide, Portlandia has broader and thinner sepals than the other eleven genera studied. They are leafy and reach their largest size, 1.4-3.5 cm. long and 0.3-1.8 cm. wide, in P. grandiflora. The sepals of P. coccinea, P. platantha, and P. harrisit are lanceolate, linear to lanceolate, and elliptic, respectively, and are 0.7—2.5 cm. long and 2—7 mm. wide The calyx lobes of many of the taxa studied bear finger-shaped out- growths on their adaxial surface near the base. Their structure is the same as that of the colleters commonly found on ane in | the Rubia- ceae, and they are discussed under the heading ‘“‘Colleters CoroLLa, Many of the flower characters discussed below are summarized in TABLE The most common corolla color among the genera is white, although several genera also have pink, red, violet, or purple flowers. The only genus in which yellow flowers occur is Schmidtottia. Of the 12 genera stud- ied, /sidorea and Schmidtottia offer both the most color variation and, to- gether with Coutaportla, the smallest flowers. In general, the larger-flowered species have funnelform corollas. Ex- ceptions are Osa, with its long, trumpet-shaped flowers, and Portlandia harristi, with large, narrowly bell-shaped ones. Coutarea is unusual among TABLE 3. Summary of flower characters in Portlandia and associated genera. COROLLA NUMBER OF LENGTH COLLETERS GENUS COROLLA COLOR COROLLA SHAPE FLORAL PARTS (cm.) FRAGRANCE ON CALYX Ceuthocar pus red funnelform 5 3-3.5 p ~ Cigarrilla white funnelform 5 4.8-7.2 + + Coutaportla white funnelform 4 2-2.5 + - Coutarea white or pink pe. base curved 5-6 (2-)5.2-7.3 + + and gibbou Cubanola white narrowly ee 5 17.8-27.5 + + Hintonia white funnelform 6,8 3-8 + — Tsidorea white, red-brown, tubular or funnelform 4-5 1.2-4 + &? +& —- rose, pale purple, or violet Osa white trumpet shaped 5) 26.5-27 . + Portlandia white, pink, or red _—_ funnelform, broadly 5 2.9-22 + & — +&— funnelform, or narrowly bell shaped Schmidtottia ochroleucous, funnelform, narrowly 5 1.4-5 ? _~ yellow, yellowish campanulate, or red, pink, red campanulate or purple Siemensia white funnelform 5 5.3-5.7 + — Tho gsennia white narrowly funnelform 5 20-26.8 ? + * Has glands on sepal edges, + = present, — = absent, ? = unknown. [6L61 VIGNVILYOd ‘OTTALV a2 JOURNAL OF THE ARNOLD ARBORETUM [voL. 60 rubiaceous plants in having zygomorphic flowers with the base of the corolla curved and gibbous. A few taxa deviate from the usual condition of 5-merous flowers. These are Coutaportla, Isidorea tetramera, and J. leonardii (all 4-merous) ; Ceuthocarpus and Coutarea (both 5- or 6-merous); and Hintonia (6- or 8-merous). Corolla length varies from 1.2 to 27.5 cm., with the smallest flowers found in /sidorea, Schmidtottia, and Coutaportia, and the largest in Port- landia grandiflora, Cubanola, Thogsenmia, and Osa. Corolla lobe aestivation for the tribe Condamineeae as a whole has been reported throughout the literature to be valvate. However, a search through published descriptions of the 12 genera included in this study yielded a variety of interpretations. Aestivation has been described for five genera: Coutaportia, narrowly imbricate (Urban, 1923); /sidorea, re- duplicate valvate (Bentham & Hooker, 1873); Portlandia, reduplicate val- vate (Bentham & Hooker, 1873), plaited-imbricative (Grisebach, 1861) ; Schmidtottia, induplicate imbricate (Urban, 1923); and Siemensia, nar- rowly quincuncially induplicate imbricate (Urban, 1923). An examination of flower buds of Portlandia, Isidorea, Cubanola, Cigar- rilla, Siemensia, Hintonia, Coutarea, Thogsennia, Schmidtottia, and Cou- taportla revealed that all have imbricate aestivation of the corolla lobes. The width of lobe overlap varies from 1 to 3 mm. In all cases where the corolla is 5-merous, the imbrication is quincuncial. Flower buds of Ceutho- carpus and Osa were not available for study. Imbricate aestivation would place Portlandia and most of the genera associated with it in the Rondele- tieae instead of the Condamineeae. Clearly, aestivation in the genera of these two tribes needs careful observation. STAMENS, STYLE, AND STIGMA. In all genera studied, the number of sta- mens is equal to the number of corolla lobes. The stamens are connate for their basal 1 to 5 mm., forming a ring that is adnate to the basal 1 to 3 mm. of the corolla. The stamens and style reach to the sinus of the corolla lobes except in Ceuthocarpus, Hintonia, and Coutarea, where they are exserted 1 to 1.5 cm. The slender filaments are villous on their basal quarter to half, except in Thogsennia, where they are villous for almost three-quarters of their length, Coutaportla, where they are villous for their entire length, and Osa, where they are glabrous. The anthers are basifixed, and dehiscence is longitudinal. The style is the same length as the stamens, plus or minus 1-2 mm. Unlike the stamen filaments, the style is glabrous. The stigmatic area consists of two (rarely three) narrow lines of papil- lae that begin at the apex of the style and continue down the outside for a distance of 0.2 to 6 cm. Usually these two lines are twisted several times around the style. There is no obvious correlation between length of the style and length of the stigmatic area. Although there is some evidence of protandry in specimens of Hintonia, 27 25 \ | , FIGURES 20-28. Flowers, stamens, aud styles: 20, Coutarea hexandra (Allen 796 (MO)); 21, Hintonia latiflora (Gentry 7064 (ps)); 22, Coutaportla ghiesbreghtiana 23, Cigarrilla mexicana (Aiello 1237 (A)); 24, Siemensia pendula; 25, Schmidtottia el- liptica (Alain 915 (NY)); 26, Schmidtottia cubensis; 27, Isidorea pedicellaris; 28, Ceuthocarpus involucratus. Bar scales = 10 [6L61 VIGNVILYOd ‘OTTAIV 54 JOURNAL OF THE ARNOLD ARBORETUM [voL. 60 the other genera appear to produce fertile stamens and receptive stigmas simultaneously. CoLLeETERS. The term ‘‘colleters” (from the Greek, meaning glued or fastened together) was first used by Hanstein (1868) to refer to secretory glands or trichomes found on bud scales, leaves, or stipules. Colleters a WH) 1 Wo | | 31 } FYI 29-31. Flowers, stamens, and styles: 29, Cubanola domingensis: ae Thogsennia lindeniana; 31, Osa pulchra (Burger 7320 (F)). Bar scales = 10m range in length from 0.2 mm. to 2 mm., and their shape varies from linear- lanceolate to ovate-lanceolate. Their structure is distinctive and consists of a central core of elongate cells with a layer of even more elongate cells perpendicular to and radiating from it. While colleters were described as early as 1849 by Weddell, we still know very little about their distribution and structural variation, and practically nothing about their secretory products or their function. Various terms have been applied to colleters. Weddell (1849), Lawson (1854), Mitra (1948), and Jayaweera (1963) referred to them as “glands,” or “glandular hairs.” Solereder (1899) and Metcalfe and Chalk (1950) called them “‘Driisenzotte” (shaggy glands). Foster (1949), Esau (1965), 1979 | AIELLO, PORTLANDIA 3D Lersten (1974, 1975), and Lersten and Curtis (1974) use Hanstein’s term “‘colleters.” Jayaweera (1963a, 1963b, 1964, 1965) used colleter number and ar- rangement as a taxonomic tool in his studies of Mussaenda. He found colleters on the calyx as well as on the stipules. During the present study also, colleters were found on both calyx and stipules (FicuREs 33-44; Taste 3). In order to examine stipules for colleters, an entire apical bud must be destroyed. Because very little material was available for study, stipules were dissected only in those taxa (Coutaportla, Cigarrilla, and Portlandia) where my own field col- lections were made, and in Siemensia pendula, where abundant herbarium material was available. All five species of Portlandia bear several rows of colleters along the base of the adaxial surface of the stipules (FicurE 41). There are no apparent differences in number or arrangement of colleters among the five species. In Cigarrilla and Siemensia the colleters are arranged in two groups of about fifteen each and are located in the lower corners of the stipules (Ficures 43, 44). Cigarrilla has a few scattered colleters toward the center of each stipule as well. In Coutaportla the entire inner surface of the stipule is covered with colleters (FrcurRE 42). They can be seen from the outside of the stipule as a fringe along its edges. Calyx colleters are found in Cigarrilla, Coutarea, Cubanola, Osa, and Thogsennia. They are also present in Portlandia grandiflora and P. harrisii, but not in P. coccinea, P. microsepala, or P. platantha. Among the four species of /sidorea examined, they were found on /. leonardii and I. pedi- cellaris, but not on J. leptantha or I. pungens (Ficures 33-40). Their structure is the same as that of the more widely known stipular colleters. Calyx colleters are absent from Ceuthocarpus, Hintonia, Schmidtottia, Siemensia, and Coutaportla. The calyx lobes of Coutaportla have red- tipped glands along the edges (F1GURE 32). There is much variation as to number and arrangement of calyx col- leters. In Jsidorea there are two per sepal, while Portlandia grandiflora has four, and Cigarrilla has six. Portlandia harrisii has a continuous ring of colleters within the base of the calyx. In Coutarea the colleters are ar- ranged in groups between the sepals. Osa, Cubanola, and Thogsennia have more abundant colleters, and these extend up onto the inner face of each sepal. POLLEN Pollen in the Rubiaceae is rather diverse. According to Erdtman (1952), 3-polycolpate, 2- to 4-colporate, 3-porate, polyforate, and nonaperturate types occur in the family. As part of the present study, pollen of the ma- jor genera of the tribe Condamineeae was examined, as was that of a 56 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 a ee a 32 if an : A My mn Ry 33 aan, 37 41 Ficures 32-44, 32, glands on edge of calyx lobe of Coutaportla ghiesbreght- tana. Bar scale = 3 . 33-40, colleters on calyx lobes: 33, Thogsennia lin- deniana; 34, Isidorea pedicellaris; 35, Cigarrilla mexicana (Aiello 1237 (a)): 36, Portlandia grandiflora; 37, Portlandia harrisii (Aiello 1306 (A)); 38, Coutarea hexandra (Allen 796 (NY)); 39, Cubanola domingensis; 40, Osa pulchra. Bar 1979 | AIELLO, PORTLANDIA S| number of genera assigned to the tribes Cinchoneae and Rondeletieae, for comparison with pollen of Portlandia. Pollen was removed from herbarium specimens and placed on aluminum stubs by means of double-stick Scotch tape. The stubs were then coated, first with carbon in two one-minute steps, and then with gold-palladium in two 1.5-minute steps, using a Technics Hummer II sputter coater. The specimens were examined with an AMR model 1000a scanning electron microscope in the secondary electron mode using accelerating voltages up to 20 KV. Polaroid type 55 P/N 4” x 5” film was used to record scan- ning electron micrographs. The micrographs were made at the Museum of Comparative Zoology, Harvard University, scanning electron micro- scope facility (funded under NSF Grant No. BMS-7412494). Acetolyzed pollen of seven genera was examined under the compound light microscope. Measurements are equatorial (width) followed by polar (length). All of the pollen studied was tricolporate, and fell into one of three main sculpture types. Type I (Ficures 45, 46). The pollen of Bikkia (14 pm. X 13 pm.), Cigarrilla (17 pm. & 25 wm.), Coutaportia (20 pm. * 20 pm.), Coutarea (20 pm. & 19 pm.), Cubanola (26 pm. &K 31 pm.), Exostema (28 pm. X 22 wm.), Hintonia (16 wm. X 24 um.), /sidorea (12 pm. X 23 pm.), Osa (26 pm. < 31 pm.), Portlandia (21 pm. X 24 pm.), Schmidtottia (29 um. < 27 pm.), Siemensia (17 pm. X 17 pm.), Thogsennia (16 pm. X 20 »m.), and the doubtfully placed Portlandia guatemalensis (17 pm. X 16 um.) is fairly uniform except for size. It is tricolporate, spherical to prolate, and diffusely foveolate, and bears echinate to bluntly echinate processes. Interpretation of SEM pictures is sometimes difficult due to swelling of the grains. Type II (Ficures 47, 48). In general, this type of pollen is tricolporate and prolate to spherical; the surface is foveolate and lacking in tectal processes. Macrocnemum (11 pm. X 19 pm.) is spherical to prolate, with large, widely scattered foveae. Rondeletia intermixta (8 ym. X 8 wm.) and R. pachyphylla (12 pm. * 14 wm.) are prolate and diffusely foveolate. Rustia formosa (15 pm. < 26 wm.) and Cinchona pubescens (15 pm. X 17 pm.) are densely foveolate. Tresanthera (18 pm. X 17 um.) is spherical and rugulate, with foveae in the depressions. Type III (Ficures 49, 50). This type of pollen is tricolporate, prolate to spherical, and foveolate-reticulate to reticulate-foveolate. Condaminea (19 pm. & 24 pm.) and Chimarrhis cymosa (14 pm. X 16 pm.) are foveolate to foveolate-reticulate. Chimarrhis latifolia (14 pm. X 17 pm.) and C. parvifolia (12 wm. X 13 pm.) are reticulate. Bathysa (9 pm. XK 16 s = 3mm. 41-44, colleters on stipules: 41, Portlandia harrisit; 42, Couta- portla ghiesbreghtiana; 43, Siemensia pendula; 44, Cigarrilla mexicana. Bar sca 2 mm. 58 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 Ficures 45-50, Pollen: 45, Isidorea brachycarpa (Carabia 3809 (GH)); Portlandia microsepala (Howard 14598 (1J)); 47, Cinchona pubescens ae 6534 (GH)); 48, Tresanthera ie (Webster 9805 (a)); 49, Pinckneya pu- nie ial 5091 (A)); 50, Rustia occidentalis (Svenson 331 (cx). Bar scales 1979 | AIELLO, PORTLANDIA 59 pm.) and Lindenia (13 um. & 14 um.) are reticulate to reticulate-foveolate. Calderonia (17 um. &K 26 um.), Picardaea (12 pm. & 22 ywm.), Pinckneya (14 um. & 25 wm.), Pogonopus (12 pm. & 15 pm.), and Rustia occiden- talis (14 wm. & 22 um.) are reticulate. As TABLE 4 shows, the Rondeletieae are not represented in pollen type TABLE 4. Distribution of pollen and wood types among genera surveyed. KoEK-NOORMAN POLLEN TYPE GENUS TRIBE* WOOD TYPE+ I Bikkia Co II I Ceuthocar pus Co I Coutaportla Ci I Coutarea Ci I Exostema Ci II I Hintonia Ci I Isidorea Co I Portlandia Co II I Schmidtottia Co I Siemensia Co I Thogsennia Co II Cinchona Ci I iBE Ladenber gia Ci i II Macrocnemum Ci I II Rondeletia R II II die ea Co I II Tresanthera Co Ill ae R I III Calderonia Co III Chimarrhis Co I III ea minea Co I Ill Linde R II III ie Co III Pinckneya Co I III Pogonopus Co I Ill Rustia occidentalis Co I * Co = Condamineeae, Ci = Cinchoneae, R = Rondeletieae. + Where known. I, and the Cinchoneae are absent from type III, while all three tribes have genera with type II pollen. The genus Rustia displays both types II and III. In addition, these two types appear to grade into one another, es- pecially when they are compared with pollen of Macrocnemum. Type II may simply be immature type ITI pollen. There is some correlation between pollen types and the wood types of Koek-Noorman and Hogeweg (1974). Future work will likely result in revision of these three tribes, since many genera appear to be misplaced. Siemensia, for example, probably belongs to the Hedyotideae rather than the Condamineeae. However, the possibility also remains that there is 60 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60 some ecological reason for the pollen differences. In general, the genera with type I pollen tend to have much larger, showier flowers than do the other genera. Perhaps these larger-flowered genera have either similar pol- linators or different pollinators with similar habits. In the case of Port- landia, for example, the white-flowered species are probably pollinated by moths, and the red-flowered species by hummingbirds. Both of these pollinators are fairly large, and both hover while feeding at the flowers. Practically nothing is known on the general subject of pollen-type/pol- linator relationships. A l — D-D-w (D-Q)-G FicureE 51. Diagrammatic cross sections of capsules showing double-walled septum: A, loculicidal splitting of capsule followed by septal splitting; B, septi- cidal splitting of capsule wall plus septal separation followed by septal splitting. B FRUITS Fruit and seed characters are very important to the understanding of the Rubiaceae, yet they have often been either underrated or neglected al- together. For example, although Baillon (1879) gave an excellent de- scription of the fruit of what we now know as Coutaportla, he did not rec- ognize the plant as a distinct genus but rather as a section of Portlandia. It remained for Urban (1923) to make a generic separation based on a number of characters including placental shape and seed attachment. Bullock (1935), in separating Hintonia from Portlandia and Coutarea, considered seeds winged vs. unwinged, and capsules septicidal vs. loculicidal as well as flattened vs. unflattened, and decided that, in spite of the fact that Hintonia has winged seeds, it is closer to Portlandia than to Coutarea. He also stated that the seeds of Hintonia are descending, while those of Coutarea are ascending, but he made no mention of seed attachment in Portlandia. When Urban (1923) separated Siemensia from Portlandia, he did so on the basis of leaf anatomy, with no discussion of differences in fruits or seeds. 1979] AIELLO, PORTLANDIA 61 The following discussion is an attempt to describe the diversity of fruit forms present among the taxa at one time included in the genus Port- andia Capsule size and shape are quite variable within, as well as among, genera and species. TABLE 5 shows length width index and length of TABLE 5, Fruit size for Portlandia and associated taxa. LENGTH X WIDTH LENGTH INDEX (cm.) TAXON 882 4.2 Cubanola domingensis 667 4.2 Thogsennia lindeniana 590 3.0 Cubanola daphnoides 411 2.6 Portlandia grandiflora 406 2.8 Coutarea spp. 378 a5 Hintonia latiflora 352 3.2 Osa pulchra 320 2.0 Portlandia harrisi 205 1.6 Portlandia platantha 192 1.6 Hintonia standleyana 178 1.6 Cigarrilla mexicana 173 1.8 Hintonia oct 170 ie Portlandia coccinea 141 1.5 Isidorea polyneura 132 1.2 Tsidorea pungens 124 1.6 Tsidorea leptantha 110 1.1 Tsidorea pedicellaris 107 see Ceuthocarpus ee 88 1.1 Isidorea brachyantha 84 12 Isidorea pata 84 1.2 Schmidtottia uliginosa 80 1,2 Schmidtottia cubensis de 0.9 eas as 66 1.1 Isidor 66 1.0 H ee ae eana 60 1.0 Schmidtottia marmorata 56 0.9 Siemensia pendul 56 0.8 Coutaportla ghiesbreghtiana 54 0.9 Schmidtottia Bee 45 0.9 Schmidtottia 40 0.8 Schmidtottia er 35 0.7 Isidorea tetramera 30 0.6 Tsidorea elliptica 25 0.5 Schmidtottia monantha 20 0.5 Tsidorea leonardi average capsules for the taxa under study. The largest capsules are found in Thogsennia lindeniana and Cubanola domingensis, while those of Osa pulchra, Cubanola daphnoides, Coutarea, Portlandia grandiflora, 62 JOURNAL OF THE ARNOLD ARBORETUM [voL. 60 and Hintonia latiflora are considered intermediate in size. Thus the two species of Cubanola are closer in capsule size to unrelated taxa than to each other. The smallest capsules are found in the smooth-leaved species of Isidorea (JI. elliptica, 1. leonardii, and I. tetramera) and in Schmidtottia monantha. The veiny-leaved species of /sidorea are in the same general size range as Siemensia, Coutaportla, Schmidtottia, Ceuthocarpus, and Cigarrilla, which are all unrelated to one another. The species of Hintonia and Portlandia have very similar capsule size ranges. Portlandia and J/s1- dorea, which are closely related genera, have no overlap in capsule size. As can be seen from TABLE 6, capsule shape is also extremely variable. TABLE 6. Fruit shape for Portlandia and associated genera. GENUS SHAPE(S) Ceuthocarpus ellipsoid to subglobose Cigarrilla obovoid or oblong-ellipsoid Coutaportla compressed ; ea ovate to circular Coutarea compressed, obova Cubanola ellipsoid, or obov cid with apex truncate ! ellipsoid to obovo Tsidorea ellipsoid-oblong e obovoid or subglobose Osa ] Portlandia ellipsoid to widely obovoid or globose, rarely ovoid Schmidtottia obovoid or ellipsoid-oblong; apex truncate Siemensia ellipsoid to obovoid; apex truncate Tho gsennia ellipsoid to oblong or somewhat obovoid The only really distinctive capsules are those of Coutaportla and Coutarea, which are highly compressed; they can, however, be distinguished from one another on the basis of size and shape. Although /sidorea species seem to fall into two distinct groups based on capsule size and leaf texture, fruit size and shape appear to have little taxonomic meaning among the genera under study beyond their use as aids in the identification of certain taxa. There are three possible processes involved in the capsule dehiscence of Portlandia and associated genera (with the exception of Thogsennia, which is indehiscent). In some genera all three of these processes occur, while in others only two are observed. The three processes are: 1) loculicidal splitting of the capsule wall, starting at the top and continuing part or all of the way to the base; 2) septicidal splitting of the capsule wall, including separation of the double wall of the septum, again starting at the top and continuing part or all of the way to the bottom; and 3 splitting of the septum walls along a mid-longitudinal line — this split- ting may begin at the top and continue part or all of the way down, or may begin at the midpoint of the septum and continue in both directions. The distinction between septal “separation” (part of process 2) and septal “splitting” (process 3) is important, and the two should not be PLACENTA SHAPE SEED ARRANGEMENT z ° 7a oO Ww w wn wn {e) a SIEMENSIA {o) c z ods se to. 58E wo fo nz o on ég° oa HORIZONTAL, PENDULOUS, OR ASCENDING z oO Ee of wo 79) Eps Shy ke WwW zu we od zea <¢ © ISIDOREA PORTLANDIA % CUBANOLA ® THOGSENNIA Too HORIZONTAL 0 5060 COUTAREA 2 y iy VERTICAL @ HINTONIA ee; BASIPETALLY BRICATE CO CIGARRILLA CEUTHOCARPUS | ACROPETALLY IMBRICATE ; OSA BASIPETAL PPRES LY SED SCHMIDTOTTIA COUTAPORTLA PENDULOUS OR ASCENDING Ficure 52. Placental shape and seed attachment and orientation in Portlandia and associated genera. [6L61 VIGNVILYWOd ‘OTIAIV JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 MUTI) TERRY a U NN cures thaOBARS 8 fae “ait ° SS 8 = = = = CA 2 = pI a a Vi 7 q Ficures 53, 54. Capsules. 53, Portlandia grandiflora (Proctor 16657 (1})): A, side view; B, top view; C, cross section, septum, placentae, seeds with per- sistent funicles shown; D, side view, wall removed to show septum, placentae, horizontal seeds with persistent funicles; E, side view, indicating loculicidal de- hiscence, septal splitting. 54, Schmidtottia elliptica: A, si i i C, cross section below middle, to show septum, seeds; D, side view, wall per- pendicular to septum removed, septum, placentae (center), and seeds shown; 1979] AIELLO, PORTLANDIA 65 E, side view, wall parallel to septum removed to show placenta, seeds; F, side view, indicating method of dehiscence (more septicidal than loculicidal) ; G, top view, to show early dehiscence (more septicidal than loculicidal); H, top view, to show late dehiscence (loculicidal and septicidal). Bar scales = 5 mm. Ficures 55, 56. Capsules. 55, /sidorea leonardii: A, side view; B, top C, cross section through middle, septum, small placentae, horizontal ak a persistent funicles shown; D, side view, indicating method of dehiscence (strong- ly loculicidal); E, - view, AS show early loculicidal dehiscence. 56, /sidorea polyneura: A, side view; B, view; C, cross section, septum, small placentae, horizontal seeds ane Mn funicles shown; D, side view, indicating method of dehiscence anaes oes E, top view, to show early loculicidal de- hiscence. Bar scales 5m 66 JOURNAL OF THE ARNOLD ARBORETUM So is — & p> ete sey ‘ Kes eis Bo Y Cy Orc SP 40'o. 's Ses, 2 F202 ON ry) CM) ANeets AWB, SSSive & si w < 4 y 5 Q 7 eS oO a n 4 2 fae ea] 79) > 2 As < 4 DM pp) a Hien ery << & WO i ow n m 2 ae 5 HO oO Qo a és gO a H VAD QA FP ou tH Z, a 23 > Zz <5 fs «8 2 A ~OR RA Fs fa sa 22 ) Q HY a a OR ee BA Dp «8 ages s) 2 ho i) a. ae ee: Z Ae < FP eae wal t2) = E a 6. 4<42% O Zz an As i} He we HP nm & v ee mH Q nn ist my owe nop &® wg So yn = Hoe HA mM a qQav oH Ow O Qe OFF AM Qa Aw ma H RS zp 26 gf S58 AS S# BS SES BS Bas BR EB GENUS aA Ge & oe he me Gh £2 Ae GE eee Be ate we 23 Ceuthocarpus l/s + A 20 Cigarrilla I/s 30 Coutaportla l/s + 4 Coutarea l/s + + + + 25 Cubanola L/s + 200 Hintonia S + B 50 [sidorea ] + + + 20 Osa S? + 10 Portlandia ] + + + 150 Schmidtottia l/s dite 14 Siemensia l/s + hs hs 60 Tho gsennia + + 200 = Character present; blank space = character absent; ? = character questionably present; | = loculicidal; s = septicidal; A = acropetally; B = basipetally. [6L61 VIGNVILYOd ‘OTIAIV | VOL. 60 JOURNAL OF THE ARNOLD ARBORETUM FIGURES 63, 64. 63, Ceuthocarpus involucratus (Shafer 8282 (Ny)). A-D, cap- sule: A, side view, to show calyx and subtending bracts; B, top view, older cap- sule, bracts and calyx worn away, early dehiscence (loculicidal and septicidal) ; C, cross section through middle, to show septum, placentae, imbricate seeds: D. side view, bracts removed, indicating method of dehiscence (loculicidal and septicidal). E, acropetally imbricated seeds in place on placenta (hidden). 64, 1979 | AIELLO, PORTLANDIA 73 Schmidtottia and Coutaportla have several characters in common, al- though they are probably not related. Both have placentae attached to the center of the septum and extending across the locules, dividing them into upper and lower portions. The flattened seeds are arranged into two groups, one in the upper portion of each locule and ascending from the placenta, the other in the lower portion and descending from the placenta. In the case of Coutaportia, the capsule is much compressed perpendicular to the septum (FicurE 60). There are usually only four seeds per locule, two in the upper portion and two in the lower. Occasionally there is a fifth seed in one of the portions. In Schmidtottia there are ten to fourteen seeds in each locule (five to seven in each portion) (Ficure 54). Usually several seeds are attached down the sides of the placentae, giving a rather imbricate appearance. However, most are above or below the placentae, and all are perpendicular to the septum and parallel to the long axis of the capsule. The capsules are not flattened. In neither Coutaportla nor Schmidtottia are there any seeds attached to the sides of the placentae opposite the septum. Placental shape and seed attachment and orientation are diagrammatical- ly summarized in FicurE 52 The distribution of important fruit and seed characters for Portlandia and associated genera is shown in TaBLe 7. On the basis of these charac- ters, genera associated with Portlandia are quite distinct from both that enus and each other. Portlandia, Isidorea, Cubanola, and Thogsennia, with their horizontal seeds, and possibly Osa as well, belong to the Con- damineeae. Based on the presence of raphides and multicellular uniseriate hairs, as well as on the numerous tiny seeds, Siemensia belongs to the Hedyotideae of the subfamily Rubioideae. Coutarea is best placed in the Cinchoneae because of its winged, vertical seeds. The positions of Hintoniza, Ceuthocarpus, Coutaportla, Cigarrilla, and Schmidtottia are uncertain. None of the above placements can be made with great certainty since a large amount of work remains to be accomplished with regard to tribal relationships within the Rubiaceae. SEEDS Seed characters have been neglected in both floras and taxonomic re- visions. This neglect is unfortunate since, in the few studies published, seed coat morphology has been shown to be helpful. A few of these stud- ies have made use of seed pictures produced by the scanning electron microscope (SEM). Such pictures will likely become a routine tool in future taxonomic studies. In the Rubiaceae, where tribal and generic characters are few and con- Cigarrilla mexicana. A-E, capsule: A, side view; B, top view; C, cross section through middle, to show septum, large placentae, imbricated seeds: D, side view, indicating metho d of dehiscence (loculicidal and septicidal) ; E, top view, to show early dehiscence (loculicidal and septicidal). F, acropetally imbricated seeds in place on spongy placenta (center), side view. Bar scales = 5 mm. 74 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 FicurEs 65, 66. Capsules. 65, Siemensia pendula: A, side view; B, top view; C, cross section through middle, septum, placentae, small seeds; D, side view, de- hiscence (septicidal and loculicidal) : E, top view, dehiscence (strongly septicidal, weakly loculicidal). 66, Osa pulchra (Burger 7320 (¥)): A, side view: B, to view; C, cross section, to show septum, placentae, seeds; D, side view, wall per- yen ndicular to septum removed, showing septum, placentae, seeds; E, side view, wall parallel to septum remov ed, to show split septum with split placenta show- ing through sht. Bar scales = 5 mm. 1979 | AIELLO, PORTLANDIA 75 troversial, seed coat characters may be of considerable value; in con- junction with other characters, they might eventually prove useful in de- tecting groups of related genera within many rubiaceous tribes. Seeds of the major genera of the tribe Condamineeae (as well as those of several genera of the tribe Cinchoneae that have in the past been as- sociated with Portlandia) were examined under the SEM for external morphological characters. For examination under the SEM, seeds were removed from herbarium specimens and treated by the same methods as was the pollen (see that section). In addition, seeds of nine species were studied both by dissec- tion and by serial sectioning. Sections made were 12 pm. thick. For the most part, the seed testa terminology used has been modified after Murley (1951). In some cases where this terminology proved in- adequate, modified pollen terminology was used. The following descriptions are the result of the SEM studies. Ceuthocarpus (FicurE 79). Seeds somewhat flattened, broadly elliptic to obovate, 3-3.33 mm. long; testa reticulate. Cigarrilla (FIGURE 80). Seeds somewhat flattened, elliptic to ovate, 3-3.4 mm. long; testa colliculate to low-tuberculate. Coutaportla (FicuRES 81, 82). Seeds flattened, broadly elliptic, 3.2-3.5 mm. long; testa low-reticulate, with a baccate tubercle at one end of each cell. Coutarea, Seeds flat, winged, elliptic, with slight indentation in wing at lateral attachment point, the central portion subcircular; testa reticulate, cells radiating from the center of the seed out to the periphery of the wing. C. alba. Seeds 8.2 mm. wide (greatest dimension), central portion 3- 3.2 mm. in diameter, wing 2.3-2.7 mm. wide laterally, 0.4 mm. wide on attachment side, and 0.8 mm. wide on side opposite attachment: testa cells mostly elongate. C. hexandra (FicurEes 87, 88). Seeds 1.2 cm. wide, central portion 3.5—4 mm. in diameter, wing 3.5—4.5 mm. wide laterally, 1 mm. wide on attachment side, and 1.8—2 mm. wide on side opposite attachment; testa cells mostly isodiametric, with wavy walls. Cubanola (Ficure 74). Seeds elliptic to broadly elliptic, 3.1-3.5 mm. long; testa reticulate to foveolate. Hintonia. Seeds flat, winged, obovate to broadly elliptic, with indenta- tion at attachment point; central portion broadly elliptic to circular; testa reticulate, cells not radiating from central point. HH: latiflora (Ficures 89, 90). Seeds 7 mm. long, central portion 2-3 mm. in diameter, wing 1. 5 mm. wide. H. octomera. Seeds 6.4 mm. long, central portion 2.2-2.9 mm. in diameter, wing 1.5—2.6 mm. wide. H. standleyana. Seeds 6.2 mm. long, central portion 1.7-2.9 mm. in diameter, wing 0.9-1.7 mm. wide. ~ 76 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 Ficures 67-72. Seeds: 67, 68, Portlandia harrisi Portlandia grandiflora (Morton 715 ( (A)); 71, 72, Isidorea brachycarpa (67, 69-71) and 5 mm, (68, 72). ae ‘Harris 11046 (us)); 69, NY)); 70, Zsidorea leptantha (Ekman 15431 (Morton 9733 (us)). Bar scales = 1 em. 1979] AIELLO, PORTLANDIA 77 Isidorea oe 70-72). Seeds similar to those of Portlandia, but 1.5-3.2 mm. long. Osa eee 73). Seeds plump, elliptic to broadly elliptic, 6.5 mm. long; testa mostly tuberculate but margins with colliculate areas; funicle (?) persistent. Portlandia (FicuRES 67-69). Seeds compressed and angulate due to pack- ing, elliptic to obovate, 2.5-3.5 mm. long; testa tuberculate with the tuberculae collapsed; funicle persistent and with cuplike swelling at apex. Schmidtottia. Seeds somewhat flattened, elliptic to ovate or obovate, 2—2.9 mm. long; testa colliculate in S. parvifolia (Ficurr 78), S. elliptica, and S. cubensis, reticulate in S. monantha, S. sessilifolia subsp. shaferi (Ficure 77), and S. uliginosa. Siemensia (F1cuRES 83, 84). Seeds broadly ovate to broadly elliptic, 0.9 m ong; testa with closely packed, deep pits, interspaces densely granulate. Thogsennia (F1icures 75, 76). Seeds broadly obovate, 2.2—2.7 mm. long; testa collapsed-colliculate to reticulate, cells elongate and wrinkled. Bik kia. B, tetrandra. Seeds broadly elliptic to circular, 1.7 mm. long; testa reticulate, cells elongate and longitudinally oriented; large, clavate tubercles around the periphery coated with waxlike particles. B. tubiflora. Seeds broadly elliptic to circular, 1.8 mm. long; testa reticulate, cells isodiametric, interspace margins granulate. B. lenormandii. Seeds broadly elliptic, 2.3 mm. long; testa reticulate, cells isodiametric. B. marianensis. Seeds obovate to obtriangular, 1.3 mm. long; testa reticulate, cells elongate and longitudinally oriented. Calderonia. Seeds very thin, elliptic with one flat side, 1.6 cm. long; testa reticulate, cells isodiametric and collapsed with the thin outer walls in various stages of adhesion to the innermost ones. Chimarrhis. Seed attachment peltate. cymosa. Seeds broadly elliptic, 0.5-0.7 mm. long; testa deeply reticulate, the cells obliquely deepened, reticulate-thickened, radially arranged on attachment side of seed and randomly arranged on other side. C, latifolia, Seeds elliptic, 0.50.7 mm. long; similar to C. cymosa except testa cells scalariform-thickened. C. turbinata. Seeds elliptic to broadly elliptic, 0.9-1.6 mm. long; similar to C. cymosa and C. latifolia except for size and testa cells, which are not as deep and are reticulate-foveate-thickened. Condaminea. Seeds narrowly obovate with truncate distal end, 0.6 m ong; testa reticulate, cells elongate and longitudinally oriented, ees spaces faintly granulate. Exostema (FIGURES 85, 86). Seeds winged, flattened, elliptic, 3.9 mm. ong, the seed portion large in proportion to wing, which varies in width from 3 mm. to 7 mm.; testa reticulate. JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 78 73, Osa pulchra (Burger 7320 (®)); 74, eis , Thog iia lindeniana (Shafer 1 4180 (us)); 78, sets mm. (73-75, 77, 78) and FIGURES 73-78. Seeds: daphnoides (Clemente 4942 (ny (ny)); 77, Schmidtottia sessilifolia var. shaferi Gah ottia jnneole (Alain 3592 (Nvy)). Bar scales = 50 um. (76). Picardaea, Seeds obovate with truncate aes oe 0.5-0.7 mm. long; testa strongly reticulate, interspaces strongly gra Pinckneya. Seeds flat, depressed-obovate, ith pe at lateral attachment 1979 | AIELLO, PORTLANDIA 79 point, 8 mm. long; testa reticulate, with tendency for cells to be ar- ranged radially from point of attachment, interspaces faintly scalariform- reticulate-thickened. Pogonopus. Seeds obovate with truncate distal end, 0.5 mm. long; testa strongly reticulate, cells somewhat elongate, interspaces faintly gran- ulate. Rustia, Seeds broadly obovate, 1.9 mm. long; testa shallowly reticulate. Tresanthera. Seeds narrowly obovate to obtriangular, 0.9-1.4 mm. long; testa reticulate, cells elongate and longitudinally oriented, interspaces sparsely granulate around margins. Within the tribe Condamineeae, seed orientation is horizontal in relation to the placenta. Except in Chimarrhis, which has peltate seeds, the attachment point on the seed is terminal. Seed shape and size and testa patterns are quite variable. On the basis of the three genera studied (Coutarea, Exostema, and Hintonia), the Cinchoneae show much variation as to seed orientation and attachment. The seeds tend . be elongate and winged and to have a reticulate testa pattern. Within the Condamineeae, Portlandia and /sidorea are closely related genera. In both, the seeds are somewhat compressed and have a tuber- culate surface. The tubercles tend to collapse during normal drying, re- sulting in some distortion of the original pattern. The funicle is persistent and bears a distinctive, basal, cuplike outgrowth that surrounds the base of the seed. The species belonging to these genera cannot be readily dis- tinguished by their seeds. n the basis of seed morphology, Cigarrilla mexicana (formerly Port- Pes mexicana), with its flattened, colliewate seeds that lack a per- sistent funicle, does not belong in Portlandia; nor does Osa pulchra (formerly Hintonia pulchra) belong in the genus Hintonia as originally described. Except for their large size, the plump, wingless seeds of Osa are very similar to those of Portlandia and Isidorea. The seeds of Hin- tonia, on the other hand, are extremely flattened, wide-winged, and finely reticulate; unlike those of Osa and the rest of the Condamineeae, they are ascending and imbricate. It is not yet clear whether the origin of the seed appendage in Osa is funicular or placental. Coutaportla ghiesbreghtiana and Siemensia pendula were originally described in Portlandia. Their seed testa patterns support Urban’s view that each represents a monotypic genus. The seeds of Coutaportla are reticulate-foveolate and are unique in having a collapsed tubercle at one end of most cells. aaa seeds are minute, and their large, deep pits are densely granulat Five of the ten haa of Schmidtottia were originally described in Portlandia. Here again, seed testa morphology supports Urban’s decision to erect a new genus. All seven species studied have reticulate-areolate to somewhat colliculate seed testa patterns that are distinct from those of other genera in the Condamineeae, with the exception of Ceuthocarpus. 80 JOURNAL OF THE ARNOLD ARBORETUM [voL. 60 Thogsennia lindeniana has been placed at times in Portlandia and at other times in a genus with Cubanola daphnoides. Its seeds are most like Figures 79-84, Seeds: 79, Ceuthocarpus involucratus (Clemente 3859 (GH)); 80, Cigarrilla mexicana (Purpus 5042 (GH)); 81, 82, Coutaportla ghiesbreghtiana (Conzatti 249 (GH)); 83, 84, Siemensia pendula (C. Wright 2677 (cH)). Bar scales = 1 mm. (79- -81), 50 wm, (82, 84), and 500 um. (83). 1979] AIELLO, PORTLANDIA 81 those of Schmidtottia but differ in being heavier and thicker and in having testa cells that are elongate, irregular, and not as completely collapsed. The seeds of Tresanthera, Condaminea, and Pogonopus are elongate- reticulate and granulate, while those of Siemensia and Picardaea are iso- diametric-reticulate and strongly granulate. Seeds of Rustia are isodia- metric-reticulate but not granulate. Bikkia shows a great deal of seed testa variation. Seeds of B. tetrandra are distinctive with their large, clavate, peripheral tubercles, as are those of B. tubiflora with their nearly circular shape and their granular inter- spaces. The other two species are distinguished from one another by the shapes of their seeds and their testa cells. Chimarrhis and Pinckneya both have thickenings within the inter- spaces of the testa cells. There the resemblance ends: Chimarrhis seeds are minute (0.5-1.6 mm.) and peltate, while those of Pimckneya are large (8 mm.) and have lateral attachment. Chimarrhis is unique in having obliquely deepened testa cells. The seeds of Calderonia are in a class by themselves. Their extreme size (1.6 cm.) and wafer thinness make them distinctive even to the un- aided eye. Seed sections indicated that in all nine cases studied, the testa pattern corresponds to the pattern of cells beneath; that is, one tubercle, or one apparent cell, coincides with one cell of the testa. In all cases the seeds were anatropous, and the embryos fell into the “axile,”’ “foliate,’ and “spatulate” categories of Martin (1946), in that the embryo is centrally lo- cated in the endosperm of the seed, the cotyledons are expanded and wider than the stalk, and the embryo is erect. See Corner (1976) and Fagerlind (1937) for discussions of seed anatomy in the Rubiaceae. Except for Osa, which has large seeds and a sizable embryo with con- spicuous venation, the genera and species investigated had embryos that were very similar in size and shape (TABLE 8) TABLE 8. Summary of three measurements of embryo size for nine taxa studied. MEASUREMENTS Length of Width of Width of TAXON embryo cotyledons hypocotyl Cigarrilla mexicana 2.04 0.52 0.28 Coutaportla ghiesbreghtiana 2.20 0.68 0.24 Cubanola daphnoides 2.20 0.60 0.24 Osa pulchra 4.80 2.00 0.44 Portlandia coccinea 1.68 0.48 0.20 Portlandia grandiflora 70 0.56 0.24 Portlandia harrisit 1.45 0.48 0.28 Schmidtottia elliptica 1.48 0.44 0.20 Thogsennia lindeniana 1.56 0.60 0.26 * All measurements are in mm. 82 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 Ne 4 FIGURES 85-90. Seeds: 85, 86, Exostema ee ee ‘(Caldwell 8773 (a). 87, 88, Coutarea hexandra (Alston 10973 (F)); 89, 90, ai latiflora (Ortega 6832 (F)). Bar scales = 1 mm. (85, 87, 89), 50 um. (86, and 500 um. (90). Certainly seed testa pattern is of some value at the generic level, but whether there is correlation between seed types and tribal characters re- 1979] AIELLO, PORTLANDIA 83 mains to be seen. The tribes Condamineeae, Cinchoneae, and Rondeletieae are in need of revision as a group. The use of such new characters as pollen and wood anatomy should help to clarify their true relationships; seed orientation and attachment may also prove useful in this endeavor. WOOD ANATOMY The wood structure of the Rubiaceae is surprisingly uniform for such a large group (Chang, 1951). In addition, transitional cell types are com- mon, often making it difficult to distinguish vessels from fibers. For these reasons, wood structure does not offer a useful set of taxonomic characters at the specific or the generic level. At the tribal level, however, wood structure has been shown to be of some value (Koek-Noorman, 1969a, 1969b, 1970, 1972). Using cluster analysis, Koek-Noorman and Hogeweg (1974) made a nu- merical taxonomic study of wood variation in the Cinchoneae, Condami- neeae, and Rondeletieae. This technique revealed two sets of highly cor- related characters in the groups studied: 1) fiber tracheids, many vessels in (short) radial multiples, parenchyma lacking — or scanty and paratrach- eal, and crystal sand present in ray cells; and 2) libriform fibers, vessels solitary, parenchyma diffuse-reticulate, and crystal sand absent. He found that each of the three tribes studied could be divided into two groups corresponding to these two sets of wood characters. A com- bined analysis of all three tribes together gave the same results: two distinct groups displaying the same two sets of characters as before. These results suggest that the key to understanding any one of the three tribes may be the simultaneous study of all three, using a number of approaches (e.g., wood anatomy, and pollen, seed, and fruit pba Two tribes might conceivably emerge in place of the present thre During the present study, the wood of four ae éf Portlandia was examined to determine the range of variation present within the genus and to enable comparison with the wood structure of the Rubiaceae as a whole. The wood of Cigarrilla was also examined. Wood samples of the other taxa under study were not available for sectioning. Each of the specimens collected included wood with bark, and voucher specimens with leaves, flowers, and fruits. It should be noted that the wood of both Portlandia harrisii and Cigarrilla was still young and con- tained pith. The wood samples and microscope slides made from them are in the Harvard Wood Collection at the Botanical Museum (Hw), and voucher specimens are in the herbarium of the Arnold Arboretum, both at Harvard University. Wood samples were cut into blocks approximately 1 cm.? and were boiled in water for one hour. The blocks were then placed in hydrofluoric acid for one to two weeks and, when soft, cut into sections 15-20 pm. thick on a sliding microtome. The sections were stained in saffranin and mounted in diaphane on glass slides. Macerations were made by boiling wood slivers in water and then placing them in a mixture of equal portions of 10 percent 84 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 60 nitric acid and 10 percent chromic acid (Jeffrey’s solution) overnight. The macerations were stained in saffranin and mounted in diaphane. In examining and describing wood structure, the following sample sizes were used: SAMPLE SIZE SAMPLE SIZE FOR THE GENUS MEASUREMENT FOR EACH SPECIES PORTLANDIA Vessel tangential diameter Zo 100 Vessel density 10 40 Vessel element length 25 100 Ray height vs 100 Ray density 10 40 Important wood measurements for the samples studied are given in BLE 9 r TABLE 9, Summary of wood measurements. MEASUREMENT Average Average vessel Average Average Vessel vessel element ray ray density diameter length density height TAXON per mm.” (um. ) (um. ) per mm. (um. ) Portlandia 288 20 677 is 472 P. coccinea var. coccinea 261 24 809 15 508 var. proctori 285 17 761 14 457 P. grandiflora 12 21 689 12 485 P. harristu 455 22 448 18 438 Cigarrilla mexicana 566 18 470 14 418 Portlandia P. Browne Growth rings. Indistinct. Wood diffuse-porous, fairly hard. Vessel elements. Mostly solitary, but some multiples and/or tangential banding may be present at beginning of growth rings; average density 288 vessels per mm.” "(range 111 to 642 vessels per mm.”, standard deviation (s) 132 vessels per mm.”); average vessel diameter 20 pm. (range 9-33 ym., Ss 6 wm.); average vessel element length 677 wm. (range 221-1104 um., s 207 »m.); perforation plates always simple; end wall slope 45°, exten- sions rarely elongate; intervascular pitting alternate, minute. Parenchyma. Apotracheal-diffuse, scanty: crystals rare, only rhombic es seen. Imperforate tracheary elements. Libriform fibers with slightly bordered pits and oblique apertures predominate. 1979] AIELLO, PORTLANDIA 85 Rays. Heterocellular; uniseriate to triseriate (in P. coccinea biseriate rays are short and have long uniseriate wings); average density 15 rays per mm. (s 3 rays per mm.); average ray height 472 pm. (range 83-1518 um., S 289 wm.); vessel-ray pitting alternate, similar to intervascular. Pith. Heterogeneous, with few large isodiametric cells, no plates or thickenings. Portlandia coccinea Sw. (description based on Aiello 1308 (A), Hw 30683). Growth rings. Indistinct. Wood diffuse-porous. Vessel elements. Mostly solitary but some multiples occasionally present; tendency for banding; average density 261 vessels per mm.” (range 199 to 332 vessels per mm.%, s 46 vessels per mm.”); average vessel diameter 24 um. (range 14-33 um., s 5.4 wm.); average vessel element length 809 ym. (range 497-1104 pm., s 151 p»m.); perforation plates simple; inter- vascular pitting alternate, minute. Parenchyma. Apotracheal-diffuse, scanty. Imperforate tracheary elements. Libriform fibers with slightly bordered pits and oblique apertures. Rays. Heterocellular with both square and upright cells, or of upright cells only; uniseriate or short biseriate with long uniseriate wings; aver- age density 15 rays per mm. (s 1.5 rays per mm.); average ray height 508 um. (range 60 pm.—1242 pm., s 337 wm.) ; vessel—ray pitting alternate, similar to intervascular; globular inclusions present. Portlandia coccinea var. proctorii Aiello (description based on Azello 1297 (A), Hw 30686). Growth rings. Indistinct. Wood diffuse-porous. Vessel elements. Solitary; average density 285 vessels per mm.” (range 243 to 354 vessels per mm.2, s 54 vessels per mm.”); average vessel di- ameter 17 pm. (range 9-28 um., s 5 wm.); average vessel element length 761 pm. (range 483-938 pm.,s 145 ym.); perforation plates simple; inter- vascular pitting alternate, minute. Parenchyma. Apotracheal-diffuse, scanty. Imperforate tracheary elements. Libriform fibers with slightly bordered pits and oblique apertures. Rays. Heterogeneous with both upright and square cells, or of upright cells only; uniseriate, biseriate, and triseriate in about equal numbers; average density 14 rays per mm. (s 2.5 rays per mm.); average ray height 457 pm. (range 166-1173 »m., s 261 p»m.); vessel-ray pitting alternate, similar to intervascular. Portlandia grandiflora L. (description based on Aiello 1304 (A), Hw 30685). Growth rings. Indistinct. Wood diffuse-porous. Vessel elements. Mostly solitary; average density 152 vessels per mm. (range 11 to 221 vessels per mm.”, s 35 vessels per mm.”); average vessel 86 JOURNAL OF THE ARNOLD ARBORETUM [voL. 60 diameter 21 pm. (range 9-33 m., s 5 wm.); average vessel element length 689 pm. (range 345-1035 pm., s 207 wm.); perforation plates simple; intervascular pitting alternate, minute. pee irs Apotracheal-diffuse, somewhat scanty to slightly banded. Imperforate tracheary elements. Libriform fibers with slightly bordered pits and oblique apertures. vs. Heterogeneous with both square and upright cells, very few pro- cumbent; mostly biseriate or uniseriate, rarely with some triseriate; aver- age density 12 rays per mm. (s 2.3 rays per mm.); average ray height 485 um. (range 41-1187 »m., s 240 pm.); vessel-ray pitting alternate, similar to intervascular. Portlandia harrisii Britton (description based on Aiello 1306 (A), Hw 30684) Growth rings. Indistinct. Wood diffuse-porous. Vessel elements. Solitary; wavy tangential banding; average density 455 vessels per mm.” (range 310 to 642 vessels per mm.”, s 128 vessels per mm.~); average vessel diameter 22 ym. (range 9-28 pm., s 12 pm.); average vessel element length 448 um. (range 221-635 um., s 102.5 um.); perforation plates simple; intervascular ee ee minute. Parenchyma, Apotracheal-diffuse, very s Imperforate tracheary elements. Libriform: ee with slightly bordered pits and oblique apertures. Rays. Heterogeneous; mostly uniseriate, a few biseriate; average densi- ty 18 rays per mm. (s 3 rays per mm.); average ray height 438 pm. (range 96-1449 ywm., s 319 wm.); vessel-ray pitting alternate, similar to intervascular. Cigarrilla mexicana (Zucc. & Martius) Aiello (description based on Aiello 1240 (A), Hw 30687). Growth rings. Indistinct. Wood diffuse-porous. Vessel elements. Solitary; occasional faint, narrow tangential banding: average density 566 vessels per mm.” (range 442 to 664 vessels per mm.’, s 66 vessels per mm.”) ; average vessel diameter 18 ym. (range 9-23.5 pm., s 3 pm.); average vessel element length 470 ym. (range 276-787 um., s 103 m.); perforation plates simple; intervascular pitting alternate, Parenchyma. Apotracheal-diffuse, scanty. Imperforate tracheary elements. Libriform fibers with slightly bordered pits and oblique apertures. Rays. Heterocellular; mostly uniseriate, very few biseriate; average density 14 rays per mm. (s 3 rays per mm.); average ray height 418 pm. (range 55-1311 pm., s 277 wm.); vessel-ray pitting alternate, similar to intervascular. Pith. Homogeneous, of isodiametric cells. Within Portlandia there is very little variation in vessel diameter and 1979] AIELLO, PORTLANDIA 87 ray density. However, there is a great deal of variation in vessel density, vessel element length, and ray height within each specimen, as well as among the species. In P. coccinea var. proctorii the rays are distinctly 1- to 3-seriate, while in the other taxa they are uniseriate or biseriate with only a few triseriate. Except for details of ray composition, the wood structure of P. coccinea var. coccinea and P. coccinea var. proctori are very similar. Cigarrilla stands apart from Portlandia in having greater vessel density, rays almost entirely uniseriate with rare short biseriate ones, and pith homogeneous and made up of isodiametric cells instead of heterogeneous. CYTOLOGY Chromosome counts have been published for approximately 55 genera in 16 tribes of the Rubiaceae. Taste 10 shows the distribution of the TABLE 10. Base chromosome numbers reported for four main subfamilies of the Rubiaceae. CINCHONOIDEAE RUBIOIDEAE IXOROIDEAE GUETTARDOIDEAE 9 9 10 10 11 11 1l 11 12 12 14 17 17 base numbers reported for the four main subfamilies (Darlington, 1955) The number ~ = 11 occurs in all four subfamilies and is also the most common base number throughout the family. The only published count for a species belonging to the tribe Con- damineeae was made by Fagerlind (1937) on Portlandia grandiflora in cultivation at Kew Gardens. His count was reported as “2n = 22?”; no explanation for his uncertainty was given. Because Portlandia flowers are both large and scarce, it is difficult to obtain flower buds at the proper stage for making chromosome counts, and, while buds were collected from four species, only one sample was collected during meiosis. That sample, from P. platantha, was taken from a plant growing near the Golden Head Beach Hotel, St. Mary Parish, Jamaica, on June 7, 1975. Flower buds were collected and preserved in Carnoy’s Fixative (1 part glacial acetic acid : 3 parts chloroform : 6 parts absolute ethanol), and herbarium specimens (Azello 1302) were made. Anthers were squashed in acetocarmine, and more than 15 separate counts were made on an anther 8 mm. long with chromosomes at metaphase II. The result was the same for all counts: eleven darkly staining chromosomes plus one lighter-staining one for a total of # = 12, a number not previously reported for the Cinchonoideae. TABLE 11. Results of test for presence of alkaloids in leaves. GENUS SPECIES SPECIMEN RESULT Ceuthocar pus involucratus (Wernham) Aiello Alain 845 (GH) - Cigarrilla mexicana (Zucc. & Martius ex DC.) Aiello Aiello 1225 (A) + Coutaportla ia ie ieee Urban Aiello 1262 (A) f+ Coutarea hexandra (Jacq.) Schum Hassler 12191 (F) ~ Cubanola daphnoides (R ee Aiello Rugel 374 (NY) f+ domingensis (Britton) Aiello Rose 4176 (NY) f+ Hintonia latiflora (Sessé & Moc. ex DC.) Bullock Waterfall 12779 (MICH ) = Gentry 5700 (MICH — lumaeana (Baillon) Bullock Steyvermark 47612a (PF) _ mera (Hemsley) Bullock Lundell 7469 (MICH) — standleyana Bullock Hinton 12004 (MICH) — I sidorea brachycarpa (Urban) Aiello Ekman 15258 (us) ~ leonardii Urban Evyerdam 56 (GH) — oblanceolata (Urban) Aiello Ekman 9588 (s) — polyneura (Urban) Aiello Ekman 15608 (BM) — Osa pulchra (D. Simpson) Aiello Burger & Liesner 7320 (F) ~ 88 WOALAXOGUV GIONAV AHL AO TVNUANOL 09 ‘T0A] Portlandia Schmidtottia Sienensia Thogsennia Portlandia (?) coccinea Sw. grandiflora L. harrisit Britton microsepala Urban platantha Britton & Harris ex Moore elliptica (Britton) Urban monantha Urban nitens (Britton) Urban parvifolia Alain sessilifolia (Britton) Urban shaferi (Standley) Urban uliginosa (Wernham) Urban pendula (Wright ex Griseb.) Urban lindeniana (A. Rich.) Aiello guatemalensis Standley Aiello 1300 (A) Aiello 1308 (A) Aiello 1307 (A) Aiello 1306 (A) Howard 14598 (1J) Aiello 1209 (A) Aiello 1302 (a) Alain 915 (NY) Alain 3442 (NY) Shafer 4450 (NY) Alain 3592 (NY) Alain 3447 (ny) Alain 3298 (Ny) Shafer 4018 (F) Leon 19049 (NY) Jiménez 4337 (US) Ekman 4689 (us) Steyermark 48889 (F) p+t++4+++ iP a Explanation of symbols: (—) negative, (f+) faintly positive, (+) clearly positive. [6L61 VIGNVILYOd ‘OTTAIV 90 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 60 OTHER ASPECTS OF PORTLANDIA BIOLOGY ALKALOIDS Alkaloids have been detected in the leaves, fruit, bark, and other parts of many genera in the Rubiaceae (Willaman & Schubert, 1961; Willaman & Li, 1970; Smolenski e¢ al., 1972, 1974, 1975). Caffeine from the seeds of Coffea spp. and quinine fiom Cinchona spp. are familiar examples of alkaloids found in the Rubiaceae. There are only two reports of alkaloids in genera within the Condamineeae (Pinckneya and Pogonopus (Fong et al., 1972; Willaman & Schubert, 1961)), and none for the genus Port- landia, although Raffauf (pers. comm., 1973) obtained a positive test with P. grandiflora. In the present study, dried leaves from herbarium specimens were tested, using the spot method described by Raffauf (1962), for presence or absence of alkaloids (see TABLE 11). This test is capable of detecting alkaloids if present at a concentration of 0.01 percent or higher. No at- tempt at qualitative analysis was made. Simple presence or absence of alkaloids, without qualitative analysis, is by itself not a meaningful taxonomic character and should not be used as the determining factor in a taxonomic decision. However, it can add support to decisions made previously on the basis of other characters. In this study, clearly positive results were obtained for only six species: Cigarrilla mexicana and all five species of Portlandia. Results were faintly positive for the two species of Cubanola, Thogsennia lindeniana, and Cou- taportla ghiesbreghtiana. The tests were negative for Ceuthocarpus, Hin- tonia, Isidorea, Osa, Schmidtottia, Siemensia, and the doubtfully placed Portlandia guatemalensis. POLLINATION Among the five species of Portlandia, P. grandiflora and P. platantha have large, white flowers, the odor of which has been described as that of Gardenia. Portlandia harrisii, with large, pink flowers, is also fragrant. Portlandia coccinea and P. microsepala have smaller, red, odorless flowers. It seems likely that the white- and pink-flowered, fragrant species are pollinated at night by hawk moths (Sphingidae), and that the red-flow- ered, odorless species are pollinated during the day by hummingbirds. However, no field observations of Portlandia pollination have been re- ported, and the question of what actually visits these species is still un- answered. The distributions of the five species of Portlandia on Jamaica are shown in Map 2. It is evident that the red- and pink-flowered species are all located inland, whereas the white-flowered species are found both inland and along the coast. Presumably these distributions are correlated with those of the pollinators, and it would be interesting to know whether similar distribution patterns occur in other plant taxa. 1979] AIELLO, PORTLANDIA 91 Siemensia, Coutarea, Cigarrilla, and Hintonia have large, white flow- ers that range in size from 5.6 to 8.7 cm. and are probably pollinated by hawk moths and perhaps also by bees. Almost certainly the extremely large, white flowers of Thogsennia, Cu- banola, and Osa, which range from 18.5 to 27.5 cm. long, are pollinated Maps 1, 2. 1, distribution of Cigarrilla (solid circles) and Coutaportla (circles with right half solid) in Mexico. 2, distribution of five species of Portlandia on Jamaica: empty circles = P. grandiflora; circles with dot in center = P. platan- tha; circles with lower half solid = P. harrisii; circles with right half solid = P. microsepala; solid circles = P. coccinea. 92 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60 by moths. Moth-pollination seems especially likely in the case of Osa, which has funnelform flowers with a narrow tube 16.5 cm. long. The remaining genera (/sidorea, Schmidtottia, Coutaportla, and Ceutho- carpus) include species with white, pink, red, purple, or yellow flowers ranging in size from 1 to 5 cm. Doubtless a great variety of pollinators, including bees, moths, and hummingbirds, visit these plants. ASSOCIATED ORGANISMS During a period of field work in Jamaica, various insects were noted in association with one or more species of Portlandia The paper wasp, Polistes crinita (Hymenoptera - Wesnidac, Polistinae), is widespread on the island of Jamaica. It constructs stalked paper nests in bushes, at approximately 1.5 meters above ground level. In St. Cather- ine Parish, individuals of Polistes crinita were observed collecting nectar from the discal nectaries of Portlandia grandiflora from which the corollas had fallen. The nectary, which is located around the base of the style, continues to secrete nectar for a day or two after the corolla and style have fallen Conotelus is a genus of sap beetles (Coleoptera - Nitidulidae), the adults of which are commonly found in flowers of Convolvulus sepium and other members of the Convolvulaceae, where they are known to feed on pollen and nectar (Nishida, 1957). Small numbers of this beetle have been col- lected in flowers of several other plant families as well, including Gardenia jasminoides of the Rubiaceae (Nishida, 1957: Boving & Rozen, 1962). In Jamaica, Conotelus was found in the flowers of three species of Port- landia (P. platantha, P. harrisii, and P. grandiflora). Capsules of Portlandia platantha and P. coccinea brought back to Cam- bridge from Jamaica contained lepidopteran larvae. These larvae made silken tubes among the seeds in the capsules and fed upon the seeds until pupation. Eclosion of the adults took place two to three months later. The adult moths were identified as members of the Pyralidae, one of the largest and taxonomically most difficult families of the Lepidoptera. Sugar ants (Paratrechina longicornis) were found collecting nectar from fresh flowers of Portlandia coccinea. Specimens of these insects have been deposited in the insect collections of the Museum of Comparative Zoology at Harvard University. In addition to these field observations, a number of associated organisms were discovered during examination of herbarium specimens. Several specimens of Portlandia, Cigarrilla, and Siemensia were noted to be covered with scale insects (Homoptera - Coccoidea). Specimens of Portlandia and Thogsennia showed damage made by an unidentified leaf miner. Leaves of Portlandia grandiflora from St. Ann Parish, Jamaica, had been attacked by leaf-cutter bees. The upper leaf surfaces of Osa pulchra from Costa Rica were covered 1979] AIELLO, PORTLANDIA 93 with a number of foliicolous lichens, and a liverwort of the family Lejeu- neaceae. An ascomycete, probably in the family Brefeldiellaceae of the Micro- thyriales, was found on the upper side of Cubanola daphnoides leaves from Cuba. Atichia millardetii, an ascomycete in the Myriangiales, was found on the upper side of the leaves of Portlandia grandiflora from St. Ann Parish, Jamaica DISTRIBUTION Distributions of the 12 genera under study are described below; those for all genera except Hintonia, Coutarea, and Osa are shown in Maps 1-6. Portlandia (5 species). Shrubs or small trees up to 6.5 meters tall. Endemic to the island of Jamaica. As shown in Map 2, the red- and pink- —20 84 a | A , 4. 3, distribution of Ceuthocarpus (circles with dot in center), Jsz- dorea Gu circles), and Thogsennia (circles with right half solid) in Cuba and Hispaniola. 4, distribution of Schmidtottia (solid circles) and Siemensia (circles with right half solid) in Cuba. 94 JOURNAL OF THE ARNOLD ARBORETUM [ VoL. 60 flowered species are found inland, while the white-flowered species are found both inland and along the coast. Portlandia grandiflora is under cultivation in southern Florida and on many West Indian islands. Isidorea (12 species). Shrubs or small trees up to 3 meters tall. En- demic to Hispaniola and eastern Cuba. Each of the species is found on only one of the two islands. Cubanola (2 species). Shrubs or small trees up to 3 meters tall. One species is endemic to the southeastern portion of the Dominican Republic, the other to eastern and western Cuba Thogsennia (1 species). Tree up to 10 meters tall. Endemic to eastern Cuba. Two plants recently found in the Dominican Republic were prob- ably introduced there. Once cultivated in Martinique. Schmidtottia (10 species). Shrubs or small trees up to 3.5 meters tall. Endemic to eastern Cuba. Ceuthocarpus (1 species). Shrub up to 4 meters tall. Endemic to eastern Cuba. Siemensia (1 species). Shrub (often pendent from cliffs) up to 2 meters tall. Endemic to western Cuba. Cigarrilla (1 species). Shrub or small tree up to 3 meters tall. Endemic to Mexico, where it is found in the states of Hidalgo, Querétaro, San Luis Potosi, and Nuevo Leon. Coutapor tla (1 species). Shrub up to 2.5 meters tall. Endemic to Mex- ico, where it is found in the states of Hidalgo, Puebla, and Oaxaca. Osa (1 species). Tree up to 15 meters tall. Known from two specimens collected on the Osa Peninsula, Costa Rica. Hintonia (4 species). Shrubs or small trees up to 4 meters tall. Found in western and southern Mexico and into Guatemala. See Bullock (1935) for list of localities. Coutarea (number of species not certain). Shrubs or small trees up to 15 meters tall. Widespread in tropical America from southern Mexico, through Central America, and into South America. Cigarrilla, Coutaportla, Cubanola, Isidorea, Portlandia, Siemensia, and Thogsennia grow on limestone, often in arid areas on rugged cliffs or along dry gullies, but also in moister places on wooded hillsides or along streams. Ceuthocarpus and Schmidtottia are found in the serpentine barrens of eastern Cuba. Osa has been collected from moist forest, but no soil type is mentioned. Soil data is scanty for Coutarea and Hintonia. Several labels mention granitic, gravelly, or sandy soil. PROPAGATION Portlandia grandiflora was introduced into cultivation in England in 1775 by a Mr. Ellis, Esq. In 1795 William Curtis wrote that, “It forms a very beautiful stove plant, not of difficult growth, and readily disposed to flower. . . .”’ According to Rees (1819), it was under cultivation in 1979] AIELLO, PORTLANDIA 95 Mapes 5, 6. 5, distribution of Cubanola daphnoides (solid circles) in Cuba. 6, distribution of C. domingensis (circles with right half solid) in the Dominican Rep ublic the Marchioness of Rockingham’s garden at Hillington. An herbarium specimen taken from a flowering plant of Portlandia grandiflora in her gar- den is in the British Museum (Natural History). Portlandia grandiflora is found in cultivation in Malaysia and on seven of the West Indian Is- lands, as well as in England and the United States. Portlandia coccinea came into cultivation in pos in 1812 (Loudon, 1832), as did P. platantha in 1850 (Hooker, 1850). Graham (1841) reported that Cubanola ae (under the name Portlandia daphnoides) was growing at the Royal Botanic Garden at Edinburgh. He compared the plant to Portlandia grandiflora and wrote, “| . notwithstanding what is said in the Bot. Mag. of the easy culture and free flowering of the P. grandiflora, it is universally acknowledged among cultivators that no plant is with greater difficulty kept in good con- dition, and scarcely any flowers less frequently.”’ The experience of the Arnold Arboretum in attempting propagation of Portlandia has been similar to that of Graham. Although one cutting brought back from Jamaica bloomed within a year, most made little progress in two years. Seeds germinated readily and formed a thick car- 96 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. 60 pet of seedlings that subsequently did nothing for months. Gradually most died, although a few managed to add several leaves. Two years later, four or five had attained a height of 20 cm. By comparison, a specimen of Portlandia coccinea grown from seed by Mr. Arthur Sutton outdoors in Jamaica became a flower-bearing shrub almost a meter tall in four years. In addition, several large shrubs of Portlandia platantha growing outdoors at the Jennings Estate in Coral Gables, Florida, blossom regularly. The key to success in propagation of Portlandia probably lies in the use of leaf-mold infected with the correct vesicular-arbuscular mycorrhiza, in- stead of sterilized soil. Vesicular-arbuscular mycorrhizae occur in associa- tion with the roots of plants in practically all families of angiosperms. There, in return for simple carbohydrates obtained from the plant, the fungus enables increased ion absorption by the plant roots. While phos- phorus is the most important of these ions, passage of nitrogen, potas- sium, calcium, magnesium, iron, boron, and manganese is also involved. The necessity of this relationship for the normal growth and development of most plants may help explain the success of horticulturists growing Portlandia out-of-doors in Jamacia and Florida, and the success of early horticulturists in England who, according to Rees (1819), Hooker (1850), and Bailey (1960), used leaf-mold, and who perhaps received Jamaican soil with their shipments of cuttings. LocaL NAMES AND USES Cigarrilla mexicana. In the state of San Luis Potosi, the colloquial name for C. mexicana is “cigarrilla” or “cigarrillo” because the leaves roll up when dry, resembling cigarette paper. A specimen collected in 1892 in Mineral de Guadalcazar (San Luis Potosi) is labeled “cacaloxochil,” an Aztec name meaning “raven-flow- er, which is white, red, or yellow and very fragrant” (Simeon, 1963). In the state of Hidalgo, C. mexicana is called “flor de San Pedro.” The people of Presa Guadalupe (Hidalgo) place the fresh leaves and twigs of C. mexicana in water and boil them. The resulting liquid is swallowed to induce vomiting. On the 15th day of May, in Santa Catarina (Hidalgo), the flowers of C. mexicana are collected and tied in bunches to decorate the church for the Feast of San Isidro. Cubanola CEPA OES, In Cuba this plant is known as “‘clarine” or “clarin de farallon. Cubanola domingensis. In Santo Domingo the name “‘campanitas” is ap- plied to a number of plants in the Convolvulaceae, as well as to C. domingensis. Isidorea leptantha and I. pungens. “Palo de cruz” is used to refer to a number of unrelated plants, in addition to these two species of /sidorea. Portlandia. On the island of Jamaica, species of Portlandia are referred 1979] AIELLO, PORTLANDIA 97 as are many other plants with opposite, shining, to as ‘“‘wild coffee,” rom green leaves. No local names or uses could be discovered either questioning the people or through examination of collection labels. Patrick Browne (1756), in his description of the genus Portlandia, gave the Aztec name “tecomaxachil” for P. grandiflora. Since it is unlikely that an Aztec name was in use in Jamaica, perhaps it was applied by a worker who thought the plant resembled a Mexican plant he had seen. ee monantha. The Cuban name for this plant is “flor de tres os 2 rae lindeniana. Cuban applied to this plant are “clarin cimarron” and “clarin del rio.’ TAXONOMY Key To GENERA PREVIOUSLY ASSOCIATED WITH PORTLANDIA BASED LARGELY ON LEAF AND FLOWER CHARACTERISTICS 1. Leaves coriaceous, somewhat thick. 2. Leaf apex mucronate or pungent. 3. Leaves oblong, apex mucronate, secondary veins 2 to 7 pairs; stipules triangular, apex acuminate to attenuate; corolla 4.8-7.2 cm. long; n 3. Leaves various in outline, apex pungent, secondary veins 10 to 3 pairs closely packed; stipules narrow with long bristle tips; corolla 1.2— 4-¢m, long; short shoots often presente wee ne seein Isidorea. 2. Leaf apex neither mucronate nor pungent. 4. Leaves, when dry, dark (dark brown to black) above = lighter (green to brown) below; stipules truncate or erose, sheathi Ds ie erose ; flowers and fruits surrounded at ae: by leafy bra ses se deiner ide eee ash ays Eden 4, heaves when dry, not strongly ene ae triangular, arate or somewhat connate at base but not sheat 6. Leaves elliptic, not shining, 2- i cm. ae .Q- 13 cm. wide, apex corolla 4-merous, 2—2.5 cm. long; fruit compressed. ...... eae Coutaportla. 6. Leaves various in outline, shining, 6-22 cm. long, 2.5-14 cm. wide, apex acute to acuminate, or rounded to obtuse; corolla 5-merous, Beare m. long; fruit not compressed. ............ Portlandia. 1. Leaves chartace thin. 7. Leaves elliptic 7 elliptic- ean 3.5-5.5 cm. long, 1.6-3 cm. wide, macu- late’, corolla ‘S-merous, 5.3—S97 Ci BIONGe i ese 58 Siemensia. ts Leaves various in outline, 4-19 cm. a eee 1.9-6.5 cm. wide, not maculate; corolla 5-, 6-, or 8-merous, (2—)3-27.5 cm. long. 8. Leaves elliptic, lanceolate, or ovate, 4-10 cm. long, 1.9-5 cm. wide; flowers 5-, 6-, or 8-merous; calyx lobes 0.7-1.5 cm. long; corolla (2-)3-8 cm. long; seeds winge 9. Flowers 1 to 3 or more per axil; corolla 5- or 6-merous, gibbous at acute; JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 60 base; fruit strongly compressed, dehiscence both ge and septicidal; seeds elliptic, eee OI SIG. rag utared. 1 per axil; corolla 6- or 8-merous; corolla not ae at fruit not strongly ee dehiscence septicidal; seeds broadly elliptic, attachment at end. ................ i Na) ry < iS a Le} 8. Leaves clipes, Da ie oe elliptic- iongechare: or ovate, 7-19 cm. long, 3.5-6.5 cm. wide; flowers 5-mero a calyx lobes 2—2.5 cm. long; corolla 17.8—27. - cm. re eee not w 10. Leaves, when dry, usually pale green, core veins 4 to 5 pairs; fruits winged; Calyx lobes Recled, cys ashdune hays _ Cubanola. Leaves, when dry, dark green or brown to almost black, secondary veins 6 to 7 pairs; fruits not winged; calyx lobes not keeled Leaves elliptic or narrowly oblong, apex attenuate; corolla trumpet shaped; fruit dehiscent... Osa. Leaves elliptic-lanceolate or oblong- lanceolate, apex acute to acuminate; corolla narrowly funnelform; fruit indehiscent. Thogsennia. eo — — — — KEY TO GENERA PREVIOUSLY ASSOCIATED WITH PORTLANDIA BASED ON CHARACTERISTICS OF FRUITS WITH SEEDS IN PLACE 1. Capsule subtended vA en folaceous bracts. | _.. Ceuthocarpus. ie oe without brac Fruit indehiscent, co with no apparent ribs; placentae (in cross sec- tion) twice forked. Thogsennia. bo Fruit dehiscent, usually ribbed (often obscurely so) or winged; placentae orked. (in cross section) not a Ww Seeds distinctly winged. 4. Capsule greatly flattened perpendicular to the septum; es small in cross section; seeds elliptic, attachment at side, 0.8-1.2 cm. long including wing, vertical, perpendicular to septum ‘and pla- centa, parallel to one another. ...................... Coutarea. 4. Capsule not flattened; placentae very large and spongy in cross section; seeds broadly elliptic, attachment at end, 0.6-0.7 cm. long including — basipetally imbricate. ............ _.. Hintonia. Seeds not winge 5. Funicle or piece of placenta persistent on seeds. 6. Seeds 5-7 mm. long, oriented ees ar the ns 8 to 10 per locule: Capsule SepliCidal, 20s... ade ax edb vs 6. Seeds 1.5-3.5 mm. long, perpendicular to the septum, many per locule; capsule strongly loculicidal. 7. Sena Is leafy; capsule broadly ribbed, acd — or smooth, dark brown to black at time of dehiscence... Mesaua sea eos Ge ee ee te ee ee oe re tlandia. 7. Sepals narrow, somewhat keeled; capsule with narrow ribs or helen beige to brown. .................. Isidorea. 5. Funicle not persistent. 8. Placentae filling middle portion of each locule and tending to divide locule into upper and lower portions; seeds flattened, vertical, perpendicular to septum, arranged with flat sides to- gether in two rows, one row suspended from placenta and lo- 1979] AIELLO, PORTLANDIA 99 cated in lower portion of el Oona erect on placenta and located in upper portion of loc 9. Capsule distinctly flattened BE to the septum; seeds 4 to 6 per locule; dehiscence almost equally loculicidal ANG ese MGl dad eka. 2s en a ata sa eee Ba we Coutaportla. 9. Capsule not especially flattened; seeds 10 to 16 per locule; dehiscence strongly septicidal, with some loculicidal split- BU: ody eee ee cpa bd Bo ee se bee Schmidtottia. 8. Placentae and seeds not so arranged. 10. Seeds tiny, 1 mm. or less across; capsule small, 1 cm. lon seater ated, AU To ns Be Aaa ees ls ase ees a a aie ma 10. Seeds 2.5-3.5 mm. oo capsules 2-6 cm. long. 11. Seeds horizontal; capsule winged; dehiscence loculici- dal - — confined mostly to the disc area of BG? Ts a ies rece veoh ete ere dae Cubanola. ba cae meat imbricate; capsule not winged; de- hiscence not confined to t the disc area. ... Cigarrilla. Portlandia P. Browne, Hist. Jamaica, 164. f. 77. 1756. TYPE SPECIES: Portlandia grandiflora L Shrubs or small trees of limestone areas, up to 6.5 meters high; all parts glabrous. Leaves sessile or with petioles up to 1 cm. long; blades entire, coriaceous, shiny, venation brochidodromous; stipules interpetiolar, widely triangular, acute at apex, somewhat connate at base, with colleters on adaxial side. Inflorescences 1- to 6-flowered, reduced cymes in upper leaf axils, perfect, homostylous, regular, 5-merous, fragrant in white- and pink- flowered species. Calyx lobes subulate to elliptic, often leafy, with colleters on the adaxial surface in some species. Corolla funnelform to broadly funnelform or narrowly campanulate, white, pink, or red, aestivation im- bricate. Stamens connate for basal 1-3 mm., adnate to basal 1-3 mm. of corolla, reaching to sinuses of corolla lobes, the filaments pubescent for basal 1.2-6.3 cm., glabrous above. Pollen tricolporate, diffusely foveolate, bearing echinate processes. Ovary 2- (rarely 3-)loculate, turbinate, the placentae linear, adnate to septum nearly its full length, the ovules ana- tropous, ca. 150 per locule; style glabrous, approximately same length as stamens, the stigmatic area of 2 (rarely 3) lines of papillae, slightly twisted around apical 0.7—6.2 cm. of style. Fruit a capsule opening loculicidally from above. Seeds perpendicular to septum, compressed and angulate due to packing, elliptic to obovate in outline, 2.5—3.5 mm. long, testa tubercu- late; funicle persistent; embryo approximately 1.6 mm. long, in center of abundant endosperm. Five species endemic to the island of Jamaica. The plant genus Portlandia is closely allied to /sidorea. In both genera the fruit is a loculicidal capsule, and the horizontal seeds have a tuberculate testa and a persistent funicle. Portlandia differs from /szdorea in that its larger, flexible, coriaceous leaves are not pungent and have few (6 to 8 pairs) secondary veins; its stipules are widely triangular, not splitting; its twigs are not grooved; and no short shoots are present. As with Cigarrilla, 100 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 60 alkaloid tests on a material produced strongly positive results in all spe- cies of Portlan S. Moore (1930) discussed the discrepancy among authors with respect to the style in Portlandia. The style has been depicted by various authors as bifid, trifid, or entire. Examination of flowers on herbarium specimens indicate that the stigmatic area consists of 2 (rarely 3 in P. grandiflora) lines of papillae slightly twisted around the apex of the style. Whether the style eventually splits into 2 (or 3) arms is a question to be answered, as Moore suggests, in field investigations. KEY TO THE SPECIES OF PORTLANDIA 1. Corolla red, odorless, 2.5-8.5 cm. long, without a true limb; calyx lobes 0.1-2.5 cm. long, 0.5-5 mm. wide. 2. Calyx lobes 1-9 mm. long, 0.5-1 mm. wide. ...... P. microsepala. 2. Calyx lobes 9-25 mm. long, 2-6 mm. wide. ............ . coccinea. 1. Corolla white or pink, fragrant, 3.5-22.5 cm. long, flaring near apex to form ae calyx lobes 0.7-3.6 cm. long, 2-18 mm. wide Corolla pink, 5-8.5 cm. long; flowers clustered in uppermost axils; leaves subsessile, oval to orbicular, apex obtuse, base cordate. ..... P. harrisii. Corolla white, often streaked with pink, 3.5-22.5 cm. ignas flowers 1 or 2 (or 3) per axil; leaves petiolate, elliptic to ovate, apex acute, base cuneate to rounde d. 4. Flowers large, 10-22.5 cm. long; calyx lobes elliptic, ee 1.4-3.6 cm. long, 3-18 mm. wide; capsule ribbed. .......... P. grandiflora. 3.5-9.9 cm. ae calyx lobes lanceolate to linear- ee. not leafy, 0.7-2.3 cm. long, 2-7 mm. wide; capsule rounded or ob- SeUee y ANGCd.. bisa cease ha esr eels hamper aes ys P. platantha. Ww aw =| ©) = oO 4 wm Portlandia grandiflora L. Syst. Nat. ed. 10. 1: 928. 1759. Type: P. Browne, Hist. Jamaica, ¢. 17. 1756, cited by Linnaeus. Portlandia grandiflora L. var. parviflora S. Moore, Jour. Bot. London 68: 108. 1930. TypE: Jamaica, without further locality, Broughton s.n. (BRIST, 1.V.). Shrub or small tree up to 6.5 meters high. Leaves with petioles 0.5—1 cm. long; blades elliptic, 7.6-22 cm. long, 4.4-13 cm. wide, acute at apex, acute to obtuse at base. Inflorescences reduced cymes in upper leaf axils; flowers 1 (or 2) per cyme, fragrant. Calyx lobes elliptic, often leafy, 1.4— 3.6 cm. long, 0.3-1.8 cm. wide; colleters present on adaxial surface. Corol- la funnelform to broadly funnelform, 10-22 cm. long, white, sometimes tinged with pink on angles, lobes 2.3-3 cm. long. Filaments pubescent for basal 3—6.3 cm. Stigmatic area 3—6.2 cm. long. Fruit ellipsoid to obovoid, 2-3 cm. long. DISTRIBUTION. Jamaica, both inland and along the coast in the parishes of Hanover, Westmorland, Trelawny, St. Elizabeth, Manchester, St. Ann, Clarendon, St. Catherine, St. Mary, and St. Andrew CIMENS EXAMINED. Jamaica. Without further locality: Armstrong s.n. (E), Macfadyen s.n. (K), Rohr s.n. (Pp), Schumacher s.n. (s), Shakespear s.n. (BM, S), 1979 | AIELLO, PORTLANDIA 101 Swartz s.n. (s), Wall in 1930 (s). Hanover: Campbellton Hill, Logwood, Adams 8653 (ucwr); near Bull Bay, Proctor 23471 (a, IJ, TEX); near mouth of Great River, Proctor 16657 (a, 1J); w. of Lucea, Adams 13213 (ucw1), Webster et 8568 (GH, IJ, MICH, $s, US); Lucea to Montego Bay, Britton 2916 (F, Green Island to Fish River Britton & Hollick 2162 (F, NY, US); Green Pe to Orange Bay, Adams 12770 (ucw1); near North Negril Point, Pr octor 23476 (a, IJ, MICH, TEX); Dolphin Head, Proctor 10053 (13), Robbins 2583 (ucwt). WESTMORLAND: Town Head to Bushmouth, Adams 12036 (UCWI). TRELAWNY: near Stewart Town, Fawcett in 1904 (BM, ucw1); Windsor estate, Proctor 15776 (tj). St. EvizaABeTH: Wallingford to Jenkins, Adams 6810 (BM, UCWI); Maggoty, Fosberg 42897 (NY); Maggoty Falls, West & Arnold 887 (Ny); be- tween Maggoty and Y.S., Clarkson & Kress 75-473 (cH); Oxford near Troy, Harris 8662 (A, F, NY, UCWI, US), 9407 (F, Ny, UcwI), Marble 715 (Ny); Kaiser mine area s. of Gutters, Howard & Proctor 13762 (A, IJ, NY, US); nw. of Alliga- tor Pond, Howard & Proctor 13813 (13); Pepper, Miller 1346 (us); Giddy Hall, Sangster in 1936 (pM, NY); Mt. Charles estate, 1 mi. ssw. of Giddy Hall P.O., Proctor (15442 (1J, US). MANCHESTER: 3 mi. sse. of Auchtembeddie P.O., Aiello ‘A); 1.5 mi. nw. of Mile Gully P.O., Proctor 22939 (13); Porus, Lloyd 1019 (¥, MO); near mouth of the Alligator Hole River, Lewis in 1954 (cH, 1J); between Green Bay and Cuckold Point, Robertson 9208 (ucw1); Gut River, Adams 6326 (BM, MO, UCWI); Long Bay, Cornman in 1965 (IJ). St. ANN: Fort Point, Discovery Bay, Asprey 2162 (ucw1), Stearn 720 (A, BH, UCWI); 2 mi. e. of Rio Bueno, Howard & Proctor 15127 (a, 1J), Proctor 35272 (13), Proctor & Stearn 11479 (13). CLARENDON: between Lluidas Vale and Croft Hill, Fosberg 42710 (Ny, US); May Pen to Chapelton, Aiello et al. 1304 (a), 1305 (A). St. CATHERINE: Rio Cobre, Britton 3091 (Ny); Bogwalk, Larter 21 (ucw1), Yuncker 18614 (ucw1); Bogwalk Rd., 4 mi. from Spanish Town, Burrowes 13025 See Lluidas Vale, Hunnewell & Griscom 14388 (GH, US); St. Clair property, 1 1 ese. of Riv enhead, Proctor 8375 (A, IJ); 1 mi. w. of Riverhead, Proctor ae B 1.3 mi. nnw. of Guanaboa Vale P.O., Proctor 30365 (IJ, NY, TEX); 2 mi. n. of Guanaboa Vale, middle of Stop and Think, Aiello 1294 (a). St. Mary: without ae: locality: Purdie in 1843 (kK). ST. ANDREW: Stony Hill, Harris 5878 (BM, UC This frequently collected, often-cultivated species is the most widespread of the genus. It is distinguished from other species by its very large, white flowers, leafy calyx, and ribbed capsules. Plants with the corolla approximately 10 cm. long have been described as Portlandia grandiflora var. parviflora, but such specimens have been collected in five parishes and seem to represent an extreme of corolla length throughout the range of the species. Since no geographic variation is involved, these specimens do not merit varietal status. Although the Broughton material, upon which this taxon was based, is missing from the Bristol Museum, there is reason to believe that it will soon be restored to its rightful place. Therefore, a neotype has not been designated here. Portlandia coccinea Swartz, Prodr. 42. 1788. Shrub up to 3 meters high. Leaves sessile or with petioles up to 5 mm. long; blades ovate to elliptic, or obovate, 6-13 cm. long, 2.5—9.8 cm. wide, acute or obtuse at apex, acuminate to acute, obtuse, or cordate at base. 12 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60 Inflorescences reduced cymes in upper leaf axils; flowers 1 to 3 per cyme. Calyx lobes elliptic, 0.9-2.5 cm. long, 2-6 mm. wide. Corolla funnelform, 2.9-8 cm. long, red with 5 white lines along the angles on inside, lobes 0.5—-1.2 cm. long. Filaments pubescent for basal 12 mm. Stigmatic area 0.7-1.2 cm. long. Fruit ellipsoid to widely obovoid, or subglobose, 1—2.1 cm. long. KEY TO THE VARIETIES Leaves ovate to broadly ovate or elliptic, apex acute to obtuse, base cordate to PSG OAC, 22a cto diine ens st peeee sense, coccimea var, coccinea Leaves elliptic to obovate, apex acute, base cuneate to attenua ens P. coccinea var. proctorii. Portlandia coccinea Swartz var. coccinea. a coccinea Swartz, Prodr. 42. 1788. Tyrer: /_ without further cality, Swartz s.n. (lectotype, s!; probable isotype, !) ere coriacea Sprengel, Syst. Veg. 1: 708. 1825 i. Leaves ovate to broadly ovate or elliptic, apex acute to obtuse, base cor- date to obtuse or acute DIsTRIBUTION. Jamaica, inland in the parishes of Westmorland, St. James, Trelawny, St. Elizabeth, and Manchester. SPECIMENS EXAMINED. Jamaica. Without further locality: Bancroft s.n. (xk), Harridle s.n. (GH), Purdie in 1844 (cH, K), Purdie s.n. (Ny), Swartz s.n. (BM, F, S). WESTMORLAND: Darliston, Orcutt 6263 (UC). St. JAMES: 1 mi. se. of Stone henge, Proctor 24713 (GH, IJ, MICH, TEX), TRELAWNY: eampoat Cave district, Howard & Proctor 14400 (a, 13, Us); Sherwood Content, Proctor 11056 (cH, IJ, NY, TEX); Barbecue Bottom, Ashton in 1965 (ucwr), Plowman 3257 (A), Powell 708 (17, NY), Webster et al. 8431 (1J, MICH, S, US); Mango Tree Hill, be- tween Burnt Hill and Spring Garden, Aiello 1308 (a), Proctor & Alain 24900 (1J, NY); Burnt Hill, Spring ae to Junction, Adams 6773 (BM, MO, Ucwr): Burnt Hill, Barkley 22/240 (13). ae ELIZABETH: Ipswich, Harris 12505 (xy, ucwI), 12515 (A, BM, CAS, GH NY, S, UCWI, US). MANCHESTER: 1.5 mi. sw. of Craig Hill P.O., Proctor ee (cH, 1). Portlandia coccinea Swartz var. proctorii Aiello, var. nov. Folia elliptica ad obovata, apice acuta, basi cuneata ad attenuatam. : Jamaica. St. Catherine, several mi. n. of Old Harbour, Proctor sR nes IJ!; isotype, TEX!). DISTRIBUTION. Jamaica, inland in St. Catherine Parish. SPECIMENS EXAMINED. Jamaica. ST. CATHERINE: near Planters Hall, Adams 10298 (ucwI); 3 mi. n. of Gutters, Proctor 35237 (13); 1.5 mi. w. of Lluidas Vale, Avello et al. 1295 (a), Proctor 28015 (13); several mi. n. of Old Harbour, Aiello et al. 1296 through 1299 (a), Proctor 32708 (1J, TEX), 32923 (13), 34233 1979] AIELLO, PORTLANDIA 103 (13), Webster & Wilson 4872 (A); 3.5 mi. n. of Colbeck, Aiello 1300 (a), 1301 (A) This variable species is distinguished from Portlandia microsepala by its larger, elliptic calyx lobes, and from the remaining species by its red flowers. Although its leaves and those of P. Aarrisii both have cordate bases, the blades differ in overall shape and size. Within P. coccinea there is a great deal of variation in leaf shape and corolla size. Corolla length appears to be mosaic throughout the range of the species, while leaf shape shows geographic variation, with specimens collected in St. Catherine Parish having the leaf base cuneate to attenuate, and those collected throughout the remainder of the range having the leaf base cordate to obtuse or acute. The differences are not sufficient to warrant definition of a new species, but are used to distinguish two varieties. Portlandia platantha Hooker, Bot. Mag. 76: ¢. 4534. 1850. Type: from a plant in cultivation at Kew, Hooker in 1850 (holotype, K!). Portlandia grandiflora var. latifolia de Candolle, Prodr. 4: 405. 1830. Type: amaica, without locality or collector (holotype, c-pc, Microfiche, IDC 800. 691: II. 1!). Portlandia albiflora Britton & Harris ex Standley, N. Am. Fl. 32: 12. 1918. Type: Jamaica. St. Andrew, Cane River Valley, Harris 9637 (holotype, NY!; isotypes, A!, BM!, Ny!, ucwr!, es Portlandia eee Britton & Harris ex S. Moore, Jour. Bot. London 68: 108. 1 : Jamaica. St. Andrew, Hall’s Delight, Harris 12669 (holotype, BM, ee a isotypes, ps!, F!, GH!, K!, Mo!, Ny!, P!, ucwr!, us!). Shrub or small tree up to 5 meters high. Leaves with petioles 2-6 mm. long; blades narrowly to widely elliptic or ovate, 8.2-20.6 cm. long, 2.7— 13.1 cm. wide, acute to acuminate or obtuse at apex, attenuate to acuminate or acute at base. Inflorescences reduced cymes in upper leaf axils; flowers 1 to 3 per cyme. Calyx lobes linear-lanceolate or narrowly elliptic, 0.7— 2.3 cm. long, 2—7 mm. wide. Corolla broadly funnelform, 3.9-9.9 cm. long, white, often tinged with pink on angles, lobes 1-1.5 cm. long. Filaments pubescent for basal 1.3-2.3 cm. Stigmatic area 0.7-1.3 cm. long. Fruit ellipsoid to widely obovoid or globose, 1—2.1 cm. long. DISTRIBUTION. Jamaica, coastal areas in the ees of St. Ann, St. Mary, Kingston, St. Andrew, Portland, and St. Thom SPECIMENS EXAMINED. Jamaica. Without further locality: Burns in 1949 (ucw1), Grabham in 1900 (F, N¥), Hooker in 1850 (xk), Robins in 1775 (BM). St. ANN: Ocho Rios, Hibiscus Lodge, Proctor 15536 (GH, 1}); Prospect, Hunne- well 18868 (13). St. Mary: Rio Nuevo, Hunnewell 15363 (GH); Boscobel, Adams 9914 (BM, Ucwr); Golden Head Beach Hotel, Aiello 1302 (a); Gibralter [Gibraltar] Beach, near Oracabessa, Proctor 7456 (a, 13); Cabarita Island, off Port Maria, Proctor 7551 (13). KIncsTon/St. ANDREW: Hall’s Delight, Harris 5598 (ucwr), 12669 (DS, F, GH, K, MO, NY, P, UCWI, US); Wareka Hill, Stewart in 1965 (ucwrI); road to W ee Haas 8922 (A, NY, UCWI), Maxon 10527 (cH, s, US), Pelton in 1956 (13), Webster 13662 (GH, 13); Long Mountain, Adams 104 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 60 6192 (ucw1), Britton 812 (NY), a kaa 13061 (vew1), Campbell 6174 (ucwh), Harris 7870 (r, NY, UCWI), 12530 (BM, CAS, GH, MO, NY, S, UCWI, US), Proctor 29165 (1J, NY, TEX), Robbins 2581 a 2582 (ucwl), Skelding 2676 (ucwl), Stearn 833 (a, BM), Weaver 916 (ucw1), Webster et al. 8369 (BM, F, GH, IJ, MICH, S, US), Vuncker 17321 (MICH, S$); Cane River Valley, Adams 6961 (BM, UCWI), Cr osby et al. 322 (F, GH, MICH, MO, NY, TEX, UC, US), duQuesnay 869 (ucwl1), Harris 9637 (A, BM, NY, UCWI, US), 12532 (A, BM, CAS, GH, MO, NY, $, UCWI, US), 12670 (BM, DS, F, GH, K, NY, S, UCWI, US); Lindo’s Gap Road, 1.9 mi. from Horse River, Aiello et al. 1289 (A), 1290 ( (a). PoRTLAND: near Hope Bay, Harris 6060 (¥, NY, UCWI); near mouth of the Rio Grande, “ IJ), 24885 (1J, MICH, TEX). St. Tuomas: Yallah’s Point, Britton 3671 (Ny); 0.5 mi. w. of White Horses, Azello et al. 1291 through 1293 (a), Proctor ee (BM, GH, IJ, MICH, NY, TEX); 2 mi. cg from Rozelle, Adams 13035 (BM, UCWI); w. of Morant Bay, Chrysler 4638 (¥ ? —~ This highly variable species is similar to Portlandia grandiflora, but the flowers are smaller, the calyx is not leafy, and the capsules are not ribbed. Portlandia platantha is the most variable species of the genus and has been known under the names P. albiflora and P. latifolia, the original epithet being ignored in the literature. Fr1GuRE 91 (map) shows the localities from which specimens named as these two species have been collected. Upon describing P. latifolia, Britton and Harris distinguished it from P. albi- flora by its “broader leaves rounded (not cuneate) at base and the calyx with longer and markedly broader lobes.” There are two main problems with this distinction. First, specimens (Webster 8369 (micu, s)) collected at the type locality for P. latifolia bear both leaf types. Second, when leaf base angle is plotted against leaf width for specimens from all known lo- calities (F1GURE 91), there is a great deal of overlap, and no clear distinc- tion can be demonstrated among populations, although there is a tendency for the leaves of plants along the northern coast to be larger and broader than those of plants along the southern coast. In addition, since corolla length varies from 5.6 to 8.5 cm. for plants labeled P. albiflora, and from 3.9 to 9.9 cm. for plants labeled P. /atifolia, it is not a distinguishing char- acter. There is no choice but to include both P. albiflora and P. latifolia under the earliest name available, P. platantha. While the type locality for P. platantha in nature is not recorded, specimens collected at the Golden Head Beach Hotel in St. Mary Parish closely resemble Hooker’s specimen in details of leaf size and shape, and corolla size and shape. Portlandia harrisii Britton, Bull. Torrey Bot. Club 39: 8. 1912. TyPE: Jamaica. Clarendon, ei Woods, Harris 10975 (holotype, NY! ; isotypes, NY!, Ucwr!, US!). Shrub or small tree up to 6 meters high. Leaves sessile; blades widely elliptic to circular, 6-14.2 cm. long, 5.7-14 cm. wide, rounded at apex, cordate at base. Inflorescences reduced cymes in upper leaf axils; flowers 3 to 6 per cyme, fragrant. Calyx lobes elliptic, 1.4-2.5 cm. long, 4-5 mm. wide; colleters present on adaxial surface. Corolla narrowly campanulate, 5-8 cm. long, pink, lobes 1.5-2 cm. long. Filaments pubescent for basal 1979] AIELLO, PORTLANDIA 105 o) O O 10 a 1) © 2 2° ° Oo ro) bet’ a0 a © © ©@ = ® © © Q % en — Oo ® e oO : é = 5 = io) e © LL e e ©