S M I T H S O N I A N C O N T R I B U T I z O N S T O B O T A N Y N U M B E R 2 9 A Study of the Tribe Gesnerieae, with a Revision of Gesneria (Gesneriaceae: Gesnerioideae) Laurence E. Skog SMITHSONIAN INSTITUTION PRESS City of Washington 1976 A B S T R A C T Skog, Laurence E. A Study of the Tribe Gesnerieae, with a Revision of Gesneria (Gesneriaceae: Gesnerioideae). Smithsonian Contributions to Botany, number 29, 182 pages, 86 figures, 9 tables, 1976.-A study is made of the tribe Gesnerieae of the family Gesneriaceae from the West Indies, giving information on history, anatomy and morphology, pollination and dispersal, and hybridization in the tribe. The tribe comprises 67 species in 3 genera: Rhytidophyllztm, Gesneria, and Pheidonocarpa. The last genus is described as new, with a new species com- bination, Pheidonocarpa corymbosa (Swartz) L. Skog, and 2 subspecies. A revi- sion of Gesneria Linnaeus is presented based on field and herbarium studies. Gesneria is divided into 9 sections, 46 species (a new species, Gesneria onycho- calyx L. Skog, is described), 12 subspecies, and 11 varieties. The taxonomic portion includes keys, synonymies, descriptions, typifications, distributions, and ecology, as well as distribution maps and illustrations of the taxa. Also enum- erated in Appendix 1 are many species names once included in Gesnera or Gesneria, but which have been transferred to other genera. Two new combina- tions are made in this portion of the text: Rhytidophyllum cumanense (Han- stein) L. Skog and Rhytidophyllum onacaense (Rusby) L. Skog. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution?s annual report, Smithsonian Year. SI PRESS NUMBER 6053. SERIES COYER DFSIGN: Leaf clearing from the Katsura tree Cercidifihyllum japonicum Siebold and Zuccarini. Library of Congress Cataloging in Publication Data Skog, Laurence E. 1943- A study of the tribe Gesnerieae, rrith a revision of Gesneria (Gesneriaceae: Gesnerioideae) (Smithsonian contributions to Botany ; no. 29) Bibliography: p. Includes index. Supt. of Docs. no.: SI 1.29:29 1. Gesneria. 2. Rhytidophyllum. 3. Pheidonocarpa. 4. Botany-Jt?est Indies. I. Title. 11. Title: Gesnerieae, with a rekision of Gesneria (Gesneriaceae: Gesnerioideae). 111. Series: Smithsonian Institution. Smithsonian contributions to Botany : no. 29. QKlS2747 no. 29 [QK495.G4] 581?.08s [583?.81] 75-61934 Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Historical Review . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pollination and Seed Dispersal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hybridization and Cytology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Systematic Treatment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tribe GESNERIEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Morphology and Anatomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key to the Genera in Tribe Gesnerieae . . . . . . . . . . . . . . . . . . . . . . . . Pheidonocarpa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gesneria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key to the Sections of Gesneria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key to the Species of Section Pentarhaphia . . . . . . . . . . . . . . . . . . . . . . . Section 2 . Stenochonanthe . . . . . . . . . . . . . . . . . . . . . . . . . Section 3 . Lachnoblaste . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Section 5 . Gesneria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Section 1 . Pentarhaphia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Section 4 . Myrmekianthe . . . . . . . . . . . . . . . . Key to the Species of Section Gesneria . . . . . . . . . . . . . . . . . . . . Section 6 . Physcophyllon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key to the Species of Section Physcophyllon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Section 7 . Dittanthera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key to the Species of Section Dittanthera . . . . . . . . . Section 8 . Duchartrea . . . . . . . . . . . . . . . . . . . . . . . . . . Section 9 . Chorisanthera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key to the Species of Section C sun thera . . . . . . . . . . . . . . . . . . . . . . Appendix 1 : Excluded Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Appendix 3: List of Exsiccatae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Appendix 2: Numerical List of Taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 1 2 6 26 33 36 36 39 40 43 45 46 46 80 85 86 90 91 100 101 121 123 137 145 145 158 165 166 171 176 ... 111 A Study of the Tribe Gesnerieae, with a Revision of Gesneria (Gesneriaceae: Gesnerioideae) Laurence E. Skog Introduction T h e Gesneriaceae, a member of lariales in the classification system the Scrophu- of Cronquist (1968), are found in tropical and subtropical regions of the world, but rarely occur wild in temperate zones. The family is composed of about 120 genera and about 1800 species; two subfami- lies are recognized, separated by Burtt (1962) on the characters of seedlings into the anisocotylous Cyrtandroideae, primarily Old World, and the isocotylous Gesnerioideae in the New World. The latter subfamily has been tentatively divided into five tribes by Wiehler (1970) using characters of epidermal cell sinuation, node vasculature, shape of the vascular crescent in the petiole, and leaf arrangement. The tribe Gesnerieae, one of the five tribes maintained by Wiehler in the Gesneri- oideae, is apparently restricted geographically to the West Indies and a few localities i n northern South America. I t is restricted morphologically and anatomically by the occurrence of alternate leaves in most of the members of the tribe, by having the vascular trace of the petiole invaginated or cylin- drical (Wiehler, 1970), by the fruticose or arborescent habit, an inferior or subinferior ovary, and by having a chromosome number of n = 7, 14, or 28. The taxonomy of all known members of the Laurence E. Skog, Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560. Gesnerieae has not been attempted since the treatments by Hanstein from 1854 to 1865. Because exploration of the American tropics in the last one hundred years has revealed many species new to science, a study of this group of flowering plants is long overdue. In the study of the systematics of Gesnerieae, perhaps the most difficult problem is generic delimitation, a problem found in other groups in tlic American Gesneriaceae (Hurtt, 1956; Moore and Lee, 1968). The factors above and those following led to the choice of the Ges- nerieae as a taxon for investigation: (1) the pos- sibility of unusual pollination mechanisms in the tribe: and (2) the availability of material of many species of the genera included in the Gesnerieae among the living collections of Gesneriaceae a t Cornell University and elsewhere. In the study of the tribe Gesnerieac and the genus Gesneria presented here, three avenues of research were explored: (1) a morphological and anatomical approach to discover or elucidate floral and vegetative characters, using living material where possible to supplement herbarium specimens; (2) a comparison of chromosome analysis with the rcsults of hybridization to suggest the genetic bases for morphological characters; and (3) field studies and investigation of possible pollination mechanisms, which may reveal the interaction of the species of the Gesnerieae with the environment and how the interaction influ- ences speciation and evolution within the group. ACKNoWLEDGMEwrS.-A substantial part of the 1 2 SMITHSONIAN CONTRIBUTIONS TO BOTANY work presented here was undertaken at Cornell University and was submitted as a thesis in partial fulfillment for the degree of Doctor of Philosophy (Skog, 1972~). I greatly appreciate the advice and assistance of the members of my special committee at Cornell University: H. E. Moore, Jr., Chairman, for his special knowledge of the Gesneriaceae; H. P. Banks, for help in problems of morphology and anatomy: and G. C. Eickwort, for advice on pol- lination mechanisms. Financial assistance from three sources permit- ted research in new areas: Horticulture Inter- change Fellowship of the Garden Club of America, the English-Speaking Union, and Fison?s Horti- culture, Ltd., allowed research at the Royal Botanic Garden, Edinburgh, during 1968-1 969; National Science Foundation Grants GB-6277 and GB-8766, to H. E. Moore, Jr., principal investigator, provided support from 1969 to 1972 and permitted a collecting trip to the West Indies in 1970; and a gift from the American Gloxinia and Gesneriad Society to the L. H. Bailey Hortorium supported the use of a scanning electron microscope to study pollen and seeds. I also am indebted to the curators of the 46 herbaria who allowed me to study their collections, to J. Wurdack and W. Dress for checking the Latin descriptions, to B. L. Burtt for offering his time and aid during my study in Edinburgh, to H. Robinson for reading the manuscript, and gratitude most of all to my wife, Judith E. Skog, for her under- standing support and help. Historical Review Charles Plumier, a French missionary, botanist, and explorer of the West Indies, published in 1703 a work about the plants of the islands he visited. In this work, Plumier named for the Swiss natural- ist Conrad Gesner a group of plants he called Gesnera. Included were three species: Gesnera humilis, Pore flavescente; Gesnera arborescens, amplo Pore jimbriato & maculoso; and Gesnera amplo digitalis folio tomentoso. Presented here is a review of the history of Plumier?s name Gesneya to the present-day genera in the Gesnerieae. During his sojourn on the island of Hispaniola in 1693, Plumier observed, drew, and described some of the plants he published in 1703. His drawings and descriptions were never published in full, although at least three sets of Plumier?s draw- ings were distributed after his death. One set was made in 1733 by Claude Aubriet for Herman Boerhaave. After the death of Boerhaave, the Aubriet copies went to Johannes Burman, who later used the drawings in preparation of Plumier?s Plantarum Americanaruin Fasciculus Primus (-Deci- mus), which he edited (1755-1760). Plumier?s original manuscript and 5081 drawings are now at the Bibliotheque Centrale of the Museum Na- tional d?Histoire Naturelle at Paris, whence copies were obtained to clarify the descriptions of Linnaeus. The Aubriet copies are now in the University Library at Groningen, Netherlands, and other sets of copies are in the Herbarium Library of the Royal Botanic Gardens, Kew, and at the Department of Botany, British Museum (Natural History), London. Gesneria humilis, G. tomentosa (= Rhytidophyllum tomentosum (Lin- naeus) Martius [in part]), and Cranz?olaria fruticosa ( = Gesneria {ruticosa (Linnaeus) 0. Kuntze), described by Linnaeus in 1753, were based on Plumier?s account in the Nova Plan- tarum Americanarum Genera, as was the genus Gesneria in Linnaeus? Genera Plantarum. Another naturalist and explorer, Hans Sloane, a contemporary of Plumier, visited British islands of the West Indies in 1687-1689. He collected speci- mens that were also used by Linnaeus as bases for his names. One of the phrase names Sloane pub- lished in 1696 and 1707 was Digitalis folio oblongo serrato, ad foliorum alas Porida later included in Linnaeus? concept of Gesneria tomentosa. Another of Sloane?s polynomials, Rapunculo afinis ano- mala vasculifera, folio oblongo, serrato, flore coccineo tubuloso, semine minuto, oblongo, luteo, for a plant from Jamaica, was the basis for Gesneria acaulis, published by Linnaeus in 1759. With the publications by Olof Swartz in 1788 and 1797-1806, nine additional species names were joined to Gesneria. By the second decade of the nineteenth century the genus Gesneria had become a repository for many gesneriaceous species from the New World, placed there by authors who inflated the generic limits set by Linnaeus to include plants from South and Central America with those of the West Indies. John Lindley was the first to try to sort out the plants included in Gesneria into more than one NUMBER 29 3 genus. I n 1827 (under pl. 1110) he wrote in Edward?s Botanical Register: T h e genus Gesneria, as it at prescnt stands, contains an assemblage of plants differing so much among each other, both in habit and parts of fructification, that it is probable a careful investigation of the species will lcad to the estah- lishment of more than one new genus. Unfortunately, Lindley proceeded to select red, tubular-flowered Brazilian species to be the ?most legitimate representatives of the genus? Gesneria, discarding Linnaeus? concept of the genus and erected two new genera, Pentarhaphia (?Penta- raphia?) and Codonophora, for the species of Swartz and Linnaeus. Pentarhaphia was typified by P. longiflora Lindley, an illegitimate renaming of Gesneria ventricosa Swartz, in allusion to its long tubular corolla. Into Codonophora he placed two distinct plants: Gesneria prasinata Ker-Gawler, renamed Codonophora grandifiora Lindley, and Gesneria tomentosa Linnaeus, which became Codonophora lanceolata Lindley. Neither species of Codonophora was selected as the type-species by Lindley, which caused Codonophora to be ignored and shifted as a generic synonym from one genus to another, depending on the disposition of the species. The problem was resolved by Hanstein (1 854), who removed Gesneria tomentosa to R h y t i d o p h y l l u m , thus lectotypifying Codonophora on G. prasinata (Morton & Denham, 1972). Gesneria prasinata is actually a species of the older genus Paliavana Vandelli, of which Codonophora Lindley is a generic synonym. While agreeing with Lindley that a large com- plex of species was assigned to Gesneria Linnaeus, Martius (1 829) disregarded Lindley?s generic names and chose three of his own. He established Gesnera (using the spelling of Plumier but ex- cluding the latter?s species as well as those of Linnaeus and Swartz) for a group of species including Codonophora lanceolata and most of Gesneria sensu Lindley. Martius disregarded Pen- tarhaphia Lindley because the name did not honor Conrad Gesner as Plumier had intended. In place of Pentarhaphia, Martius named a new genus, Conradia, to include Gesneria humi l i s Linnaeus and Gesneria craniolaria, G . scabra, G . ventricosa, G. exserta, G . calycina, and G. p u m i l a of Swartz. Conradia, as well as being illegitimate when named by Martius, was also a later homonym of another generic name, Conradia Rafinesque, published in 1825. Conradia was subsequently chosen illegiti- mately as a generic name by two later authors: Nuttall (1834) and Kufferath (1914). Other species included earlier in the genus Gesneria Linnaeus-G. tomentosa L. and G. grandis Swartz-were placed in a new genus R h y t i d o p h y l l u m by Martius, which also included a new species R. berteroanum. During the sixty years following the description of the new genera Gesnera, Conradia, and R h y t i - d o p h y l l u m by Martius, many additional species of Gesneriaceae from the New World were named. During this time, however, new genera also were described, which absorbed many of the species included in Gesnera Martius. Thus, by 1890, the concept of Gesnera had nearly disappeared except for the occasional usage in horticulture and in popular journals. Appendix 1, a list of excluded species, gives names that have been included in Gesnera Martius (Ge.meria of auct., non Linnaeus) and Gesneria Linnaeus, which now best reside elsewhere. Authors following Martius a t first accepted his disposition of the species. George Don (1838: 650-651) divided the species that Martius had included in Conradia into three sections distin- guished by the relative exsertion of the stamens and coherence of the anthers: Sect. I. Synanthera. . . . Stamens enclosed. Anthers all Sect. 11. Dittanthera. . . . Stamens exserted; anthers con- Sect. 111. Chorisanthera. , , . Stamens a little exserted; connected into a round head. nected by pairs. anthers distinct. Fruit likc those of the other species. Don (1838) also erected the tribe Gesnerieae, but included in it many genera later transferred to other tribes. His subtribe Conradieae is now equivalent to the present tribe Gesnerieae. Shortly after the publication by Don of Ges- nerieae, Endlicher (1836-1 850) described Euges- nereae, roughly synonymous with Gesnerieae G. Don. Eugesnereae described by Hanstein in 1854 refers to a group of plants distinct from those included by Endlicher. A. P. de Candolle (1839) adopted the name Gesnerieae G. Don, but with a slightly different circumscription. However, de Candolle accepted the sectional alignments of Don and named two new species, Conradia pedunczilosa (= Gesneria pedunculosa (A. P. de Candolle) Fritsch) and C. 4 SMITHSONIAN CONTRIBUTIONS T O BOTANY cuneifolia (=Gesneria cuneifolia (A, P. de Can- tlolle) Fritsch), based on plants collected in Puerto Rico by Sesse and Mocino. He also renamed Gesneria acaulis Linnaeus as Conradia sloanei .A. P. de Candolle. I n 1846, Decaisne wrote a monograph of the genus Pentarhaphia, adopting Lindley?s generic name for the section Dittanthera G. Don, which, according to de Candolle, included shrubs with flowers having stamens exserted and the anthers coherent in pairs. Decaisne (1846:98) placed 15 species into Pentarhaphia by erecting new species and splitting others, such as Gesneria uentricosa Swartz, because the ?meme nom specifique eait assigne a des plantes completement distinctes par la forme et la couleur des flcurs.? He also included the description of a new genus, Duchartrea, based on collections from Cuba by Linden. The separation of the shrubby species of Gesneria Linnaeus into Pentarhaphia Lindley by Dccaisne thus forced other authors to consider the remaining species as belonging to Conradia, a practice followed for forty years. The assignment of the species in this manner appears to have caused many botanists to be confused about the generic limits. W. Hooker (1851:pl. 874) wrote following his description of Conradia neglecta (= Gesneria p u m i l a subsp. neglecta (W. Hooker) L. Skog): ?This plant belongs, I presume, to the true Conradiae, not to the first division or groupe [sic] in De Candolle, which constitutes the genus Pentaraphia of Dr. Lindley; but the distinguishing characters of which I by no means understand.? Decaisne (in Herincq, 1848) described another new genus, Herincquia , in which was placed a species he believed had been assigned incorrectly. He wrote that the species did not belong in Gesneria, where it had been described in October 1846 as Gesneria libanensis by Morren, or in R h y t i d o p h y l l u m , in which it had been described in December 1846 as R. f i loribundum b y Lemaire. Evidently, Hanstein was not aware of Decaisne?s genus Herincquia , for he published in 1854 another new generic name based on the same species, O p h i a n t h e libanensis (Linden ex Morren) Hanstein. In Centralamericas Gesneraceer, published in 1858 and reprinted in 1861 by Andcrs grsted, two new genera were split off from the Conradia- P e n tarhaph ia complex. Section C h orisan t h era of G. Don was raised to generic rank for the species resembling Gesneria p u m i l a , plants with very short stems and campanulate corollas. Orsted ap- parently described the new genus based on the description of Conradia p u m i l a (Swartz) Martius in G. Don?s 1838 publication. Included in the genus were three species: Chorisanthera p u m i l a (Swartz) Orstcd; C. hispida (Bentham) Orsted, transferred from Conradia; and C. tenera Orsted, described from an Orsted collection ostensibly from Costa Kica, but which so resembles Gesneria p u m i l a Swartz from the description that one is led to believe tliat Orsted collected Gesneria p u m i l n during his 1845 sojourn in Jamaica and then accidentally mixed this material with plants from Costa Rica. There is no known type-specimen of Chorisanthera tenera in the Orsted herbarium at Copenhagen (C) or elsewhere. Orsted apparently did not realize that the name Chorisanthera had been used in the generic sense in the Gesneriaceae by Herincq in 1852. Herincq?s use of Chorisanthera for some cultivated plants of uncertain origin is not valid because he did not provide a generic description, and he cited an earlier name as a synonym under the species (Skog, in press). Another genus circumscribed by Orsted, Codon- oraphia, was to include the species of Pentarhaphia with campanulate corollas, restricting Pentarha- p h i a to those species with a narrowed tubular corolla. T h e six combinations Orsted made in Codonoraphia have not been recognized by many authors nor picked up by the editors of I n d e x Ke w ens is. Orsted (1858) also proposed new tribal and subtribal names. Part of his tribe Rhytidophylleae subtribe Eurhytidophylleae is now assignable to the present-day concept of Gesnerieae. Grisebach maintained Conradia and Penta- rhaphia in his works (1860-1862, 1866) dealing with the collections of Charles Wright from Cuba. But in his Flora of t h e British West I n d i a n Is- lands, published 1859-1 864, Grisebach described another genus, Vaupellia, for a single species formerly placed in Gesneria by Swartz and Conradia by Martius, Vaupel l ia calycinn (Swartz) Grisebach. Hanstein (1854-1865) was the first author to treat all of the species in Conradia and Penta- rhapia, as well as in the restricted genera Duchartrcn, Her incquia , Ophianthe , and Choris- NUMBER 29 5 anthem as members of one genus. In his monu- mental study of the Gesneriaceae, Hanstein (1865: 292-305) submerged all of the names under Pentarhaphia Lindley, which he divided into four unnamed sections, and a group of ?Species inter Pentarhaphiam et Rhytidophyllum dubiae.? The sections were defined on the basis of the presence or absence of costae on the calyx (or more cor- rectly, floral tube) and on the shape of the corolla: Sectio 1. Calyx vix costatus, Corolla elongata-tubulosa. An- nulus quinquelobus. Sectio 2. Calyx costatus. Corolla tubulosa. Annulus integer. Sectio 3. Calyx costatus. Corolla obliquc subcampanulata v. cyathomorpha. Sectio 4. Calyx non costatus. Corolla oblique cyathomorpha. Bentham (1876) in his treatment of the Ges- neriaceae followed Hanstein in combining the complex into one genus. Although Bentham real- ized that the generic name Gesneria Linnaeus was an older name, he (1876:1005) wrote: ?Nomen Linneanum Gesneriae strict0 jure prioritatis ad hoc genus retinendum foret, sed nomen Gesnerae, Mart., ab omnibus botanicis et hortulanis ad genus Brasilianum addictum, hodie non nisi maximo in- commode mutaretur.? Bentham (p. 1005) proceeded to divide the genus Pentarhaphia into five sections in the man- ner of Hanstein, but supplied names to these sections: 1. Eupentarhaphia. Frutices glabri, ramosi, resinosi. Pedun- culi axillarcs, elongati, 1-flori v. 3-chotome -flori. Corolla incurvo-tubulosa, parum ventricosa. Stamina exserta. This section was based on the concept of Penta- rhaphia used by Grisebach, but for plants mostly from Cuba. 2. Duchartrea. Frutex glaber, rigidus. Pcdunculi axillares, elongati, apice m-flori. Calyces ecostati, insigniter verrucoso- rugosi. This section was based on thc genus Duchartrea Decaisne. 3. Codonorafihia. Frutex glabcr. Pedunculi axillarcs, elongati, 1-flori. Corolla latc campanulata, staminibus cx- sertis. Section Codonoraphia was based on Orsted?s genus of the same name. 4. Vaupellia. Habitus fere Synantherae. Corolla ct stamina Codonoruphiae. Calycis limbus amplus, lobis latis base con- natis. Vaupellia calycina (Swartz) Grisebach was the basis for this section. 5. Synanthera, Don. Caulis humilis, simplex v. decumbens, subramosus, rarius elongatus. Pcdicelli breves, fasciculati, rarius clongati I-flori. Corolla tubulosa rarius campanulata v. ventricosa, staminibus inclusis v. corollam non super- antibus. This section comprised most of the species known at that time. The concept of the section Synanthera equals Conradia Martius as used by Grisebach in Flora of the British West Indian Islands (1859- 1864) and included Ophianthe Hanstein and Herincquia necaisne (misspelled Henrincquia by Bentham). Chorisanthera Orsted was included in Pentarhaphia, but not placed in a particular sec- tion. Bentham (1876) also described a new sub- tribe, Pentarhaphieae, equivalent to the present tribe Gesnerieae. Otto Kuntze (1 891 :473) criticized Rentham?s decision to maintain Gesnera Martius and Penta- rhaphia Lindley: BHgp. [Bentham in Bentham & Hooker?s Genera Plantarum] sind sich daruber auch klar gcwesen, stellten abcr den rechtmassigen Namen aus Bequemlichkeit nicht wicder her, licsscn also Gesnera Martius gelten, wahrend sie doch incon- sequcntcrweise den fruhcr (cfr. DC. prodr.) ebenso gebrauch- lichcn Namen Conradia Martius durch den von Lindley gegebenen Pentaraphia ersetzen. Kuntze (1 891) resurrected Gesneria Linnaeus, but preferred the spelling Gesnera as proposed by Plumier. Under Gesnera Linnaeus he submerged Pentarhaphia Lindley, Conradia Martius, and even included Rhytidophyllum Martius. Rhy- tidophyllurn Martius was embraced because Kuntze (1891:473) believed and wrote: ?Die letzteren 2 Gattungen [Conradia Martius and Rhytidophyl- l u m Martius] sind so schwach getrennt, dass BHgp. [Bentham in Bentham & Hooker?s Genera Plan- tarum] selbst meinen, es sei besser, sie zu vereinigen.? Kuntze listed and transferred many names to Gesnera Linnaeus. Three years later, in 1894, Fritsch completed the account of the Gesneriaceae for Engler & Prantl?s Die natiulichen Pflanzenfamilien. He used the name Gesneria Linnaeus, but excluded from i t Rhytidophyllum, which Fritsch con- sidered a distinct genus. His comments (1 894: 184) provide a good summary of the state of the tribe Gesnerieae at his time: 6 SMITHSONIAN CONTRIBUTIONS TO BOTANY Die ganze Gruppe der Gesnerieae ist, namentlich in Bezug auf die Ausbildung der B1. [Blute], sehr formenreich, und es sind daher mehrere Gattungen innerhalb derselben unter- schieden worden. Die beste derselben ist die Gattung Rhytid- ophyllurn, welche auch habitue11 gut charakterisiert ist; die ubrigen mussen, so auffallend sie in typischen Arten sind, wegen der zahlreichen Mittel formen als Sectionen von Gesneria aufgefasst werden. Fritsch divided Gesneria into five sections follow- ing Bentham (1876). His ?Section I? was called Pentarhaphia, differing slightly from Bentham?s Eupentarhaphia, and Fritsch?s fifth section was Conradia, apparently rejecting the earlier and validly published sectional names Synanthera and Chorisanthera of G. Don (1838). The tribe Gesnerieae was again erected by Fritsch (1894), although he does not attribute the name to G. Don, who proposed it in 1838. Fritsch?s circumscription of the tribe included only Gesneria Linnaeus and Rhytidophyllum Martius. In January of 1900, there appeared the first mention of Gesneria by Ignatz Urban, who in the next 30 years was to describe more new species of Gesneria than any other botanist. In the second volume of Urban?s Symbolae Antillanae (1901), the chapter entitled ?Enumeratio Gesneriacearum? contained an account of all the species known at that time to grow in West Indies. Urban did not divide the genus into sections, but in his enumera- tion and discussion of the species he followed the sequence set by Bentham (1876) and Fritsch (1 894). Urban described several new species based on collections of Fuertes, Harris, Shafer, and others in subsequent volumes of Symbolae Antil- Zanae (1 898-1 928). Beginning in 1914, Urban began a collaboration with Erik Ekman, the Swedish plant collector who made extensive collections on the islands of Cuba and Hispaniola. Urban based many new species of Gesneria on Ekman collections. Fortunately for later students of West Indian botany, the Ekman collections selected as holotypes were deposited in the Natur- historiska Riksmuseum in Stockholm instead of at Berlin. Because much of the herbarium at Berlin was destroyed during the Second World War, many of Urban?s names typified by collections at Berlin must now be interpreted on the basis of other material. The most recent works were papers by C. V. Morton (1957a, 1957b), who wrote the account of Gesneriaceae for the Flora de Cuba; Alain Liogier, who described a number of new species of Gesneria from the Dominican Republic (1968, 1971a, 1971b, 1973); Wiehler (1971), who trans- ferred species from Cuba and Jamaica formerly in Heppiella to Gesneria; and Skog (1972a), who published new species of Gesneria from Haiti and Jamaica. The species transferred by Wiehler in the writer?s opinion represent a distinct genus Pheidonocarpa, described later. Throughout the history of the tribe Gesnerieae, most of the works have been taxonomic. Papers dealing with anatomy and morphology (e.g., Sachs, 1915) have included some details of the plants in this tribe. Cytological results have been reported in many papers by Lee (1962, 1964, 1966a, 1966b, 1968) and others. Pollination studies by Vogel (1969a, 1969b) have revealed new mechanisms possible in the tribe. An elaboration of these topics will follow later in this paper. Morphology and Anatomy The morphology and anatomy of the tribe Gesnerieae have been examined by several investi- gators. Wiehler (1970) dealt with many of the characters in comparing the tribes of the subfamily Gesnerioideae. Sachs (1915) investigated system- atically the leaf anatomy of many members of the Gesnerieae in her report on the subfamily. The anatomical works by Solereder (1908) and Met- calfe and Chalk (1950) included characters useful in defining taxa. Some aspects of the morphology and anatomy of the tribe Gesnerieae, primarily the genus Gesneria, examined in the field and in the laboratory, are considered in an attempt to relate and interpret the anatomy and morphology to the systematics of the group. OBSERVATIONS HABIT A N D HABITAT.-Plants in the tribe Ges- nerieae are perennial, producing flushes of growth and flowers at various times of the year. Depending on the species, plants can be (1) acaulescent herbs, e.g., Gesneria reticulata (Figure Ig) , G. pumila subsp. neglecta (Figure le); (2) suffruticose with short qrect or decumbent woody stems, e.g, NUMBER 29 7 Pheidonocarpa corymbosa (Figure Id); (3) tall spreading shrubs, e.g., G. alpina; or (4) arbores- cent up to 10 m tall, e.g., Rhytidophyllum grande. T h e stems may or may not branch from axillary buds, resulting in a columnar growth in un- branched species, e.g., Rhytidophyllum grande (Figure 1 c), Gesneria viridipora subsp. quis- queyana. The taller species-Gesneria exserta, G. calycosa, among others-frequently branch at ground level. Acaulescent herbs occur often on ledges in cliff faces; these same species when grow- ing on sloping banks or cliff faces become pendent or decumbent, e.g., G. citrina. T h e habitats of the species may affect the gross morphology, particularly height, size and shape of leaves, and size and color of flower in a single population, Populations of two species, G. humilis (Figure l a ) and G. paucipora, growing in open exposed, drier areas, appear morphologically simi- lar to those frequently inundated for short periods of time in streams swollen by tropical rains; the latter populations are characterized by apparently stunted stems, reduced leaves and inflorescences. They may be functional xerophytes. Room-Fibrous roots, which may become quite large and woody with age, are found in all species of this tribe. Adventitious roots may be formed from nodes on intact plants growing in the very humid atmosphere of a greenhouse or on stream banks or in the spray of waterfalls-e.g., G. humilis, G. reticulata, G. paucipora, G. viridipora subsp. quisqueyana. The acaulescent species and those with short stems develop roots from cuttings in a short time. The taller species-e.g., G. caly- cosa, G. exserta, Rhytidophyllum tomentosum-do not root easily from green cuttings. Even when dusted with rooting hormone, two to four months may be required before a callus forms and roots appear. STEMS.-A~~ members of the tribe have woody stems, at least at the base of the plant. The bark on older stems is rugose, verrucose, or smooth. Lenticels are commonly very elongated and erumpent. The sapwood is green to white, or occasionally pink; the heartwood and pith are usually pink to red. The red color is probably due to the presence of 3-desoxyanthocyanidins found also in the leaves and flowers of most members of the subfamily Gesnerioideae (Harborne, 1966). In Gesneria three types of 3-desoxyanthocyanidins have been identified: gesnerin and luteolinidin 5-glucoside in Gesneria cuneifolia and pelargoni- din 3-rutinoside in G. uentricosa (Harborne, 1966). The pith of young stems is composed of thin- walled parenchyma cells usually containing chloro- plasts. Here, too, are found crystals, which may be calcium oxalate. Sachs (1915) stated that, until the time of her studies, raphides were unknown in the Gesneriaceae; she found them subsequently . in many genera. The neeclle-shaped raphides are found in nearly all the species of Gesnel-ia investi- gated. One exception is G. paucifiora, which has prismatic styloid crystals like those common in Rhytidophyllum and Pheidonocarpa. The stems of Gesneria ventricosa (as Penta- rhnphia Zongipora-Solereder, 1908) and Rhytido- phyllum tomentosum (Pomrencke, 1892) show the following characters: (1) the wood with vascular rays of varying widths; (2) vessels of different diameters with the maximum between 0.03 and 0.108 mm; (3) vessel perforations simple, walls with bordered pits; (4) lumina of the vessel ele- ments septate; (5) cork cells with thin walls; (6) outer portion of the primary cortex collenchyma- tous; (7) outer phloem with groups of elongated and pitted, yellow-walled, sclerenchymatous cells resembling sclereids; and (8) the phloem strength- ened by layers of suberized cells formed externally to the phloem from the primary cortical paren- chyma. Douliot (1889) found that the cork develops internally to the sclerenchyma of the outer phloem in Gesneria (as Pentarhaphia) and Rhytidophyllum plumerianum. Metcalfe and Chalk (1950) report that vessels are formed in groups of four or more cells and produce a radial pattern. The investigation of nodal anatomy by Wiehler (1970) has shown that nodes in the members of the tribe Gesnerieae are unilacunar with one trace leading to the leaf. A reexamination showed this to be true. LEAvES.-The dorsiventral leaves are spirally arranged in a 2/5 phyllotaxy in all members of the tribe, except for an aberrant form of G. fruti- cosa (L. H. Bailey Hortorium, Cornell University accession number G-1035) grown from seed in the greenhouse. These latter plants have the leaves arranged in an opposite decussate pattern, rather than alternate. Comparison of the nodal structure of the aberrant form of G. fruticosa (Figure 2b) with that of Pheidonocarpa col-ymbosa was one of 8 SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 29 9 the points of evidence used by Wiehler (1971) to unite this latter taxon, formerly in the South American genus Heppiella, with Gesneria. Siblings of the aberrant plant, however, have alternate leaves (R. Katzenberger, in litt.), although speci- mens of the seed parent show an opposite arrange- ment (Talpey 71, BH). Collections of Gesneria fruticosa from the same locality in the Dominican Republic by Ekman in 1929 (Ekman H11334) show examples of opposite leaves. Efforts to locate the plants in the wild in 1970 failed to turn up the aberrant form. All other collections of G. fruticosa observed in the wild or as specimens from many herbaria have alternate leaves (Figures 2a, 75a). T h e alternate arrangement of leaves in Gesneria and Rhytidophyllum is not common among other members of the Gesneriaceae. In other genera of the family where the arrangement is found, it is believed that alternate leaves are the result of reduction of one of the pairs of opposite leaves (Fritsch, 1893-4) based on the occurrence of an intermediate stage in which the opposite leaves are unequal. Since leaves of Gesneria and Rhytid- ophyllum are occasionally approximate in slowly growing plants, an alternative suggestion may be that the alternate leaf arrangement is due to suc- cessive plastochrons of unequal duration in periods of rapid growth (Esau, 1965). Even though the arrangement of leaves in the aberrant Gesneria fruticosa appear opposite, a transverse section at the node (Figure 2b) shows the leaves in not exactly the same stage of development. If Pheidon- ocarpa corymbosa is truly a member of this tribe, as I believe, it may considered more primi- tive in leaf arrangement than Gesneria or Rhytidophyllum. FIGURE 1.-a, Habitat and presumed type locality of Gesneria humil is at Fond de Baudin, Haiti (arrow = plants at water level); b, habitat of Rhytidophyl lurn leuconal lon in Haiti; c, habitat of R. grande in Jamaica (peduncles ca 0.5 m long); d , habitat of Pheidonocarpa corymbosa in Jamaica; e, habit of Gesneria purnila subsp. neglecta, G-875; f , habitat of C. pedicellaris on limestone cliff face in the Dominican Republic (arrow indicates horizontally oriented flower); g, habitat of G. reticulata near waterfall in Pucrto Rico; h, flower of G. fruticosa, G-1035 (scale line = 1 cm); i, infloresccnces of G . pedunculosa (scale line = 5 cm); j , flowcrs of putative natural hybrid and presumcd parent species (scale line = 1 cm); top, G. acaulis, G-S76; middle, putative natural hybrid of G. acaulis and Rhyt idophyl lum tomentosum, (3-1371; bottom, R. tomentosum, G-841. Stipules have never been reported in any mem- bers of the Gesneriaceae. Cronquist (1968) wrote that the Gesneriaceae as others in the Scrophu- lariales ?uniformly lack stipules.? Stipule-like structures, however, are evident on many plants of Rhytidophyllum, and have been discussed by Weber (1 973). These ?pseudo-stipules? or ?vagi- nalen Ohrchen? in the sense of Weberling appear in Rhytidophyllum tomentosum as reniform or auriculate flaps of tissue on the stem at each side of the petiole base or, in some examples in R. auriculatum, as lobes laterally adnate to the petiole and resembling segments of a compound leaf. The development of the ?pseudo-stipules? can be observed on Rhytidophyllum auriculatum in the greenhouse. A series of leaves borne on one axis shows the following stages (Figure 3): a petiolate leaf lacking stipules, a leaf with a narrowly decurrent base, an auriculate leaf decur- rent on one side of the petiole, and a leaf with ?pseudo-stipules? and a petiole. Apparently the a I 5 mm I FIGURE 2.-Nodes of Gesneria fruticosa: a, plant G-1379, indi- cating usual altcrnate leaf arrangement; b, plant G-1035, showing aberrant opposite leaf arrangemcnt (after IViehlcr, 1970, 1971). 10 SMITHSONIAN CONTRIBUTIONS T O BOTANY ?pseudo-stipules? are remnants of the auriculate leaf, which has regained a definite petiole by loss of the decurrent leaf margins. ?Pseudo-stipules? may appear seasonally in response to differing ecological conditions or growth rates. In any one species, however, not all populations show the stipule-like structures. In Gesneria no such elaborate structures as ?pseudo-stipules? have been found, but ridges may be present in some species, which may continue up the stem for a short distance from the petiole base (e.g., G. humilis), or protuberances may be present on the stem at either side of the petiole bases (e.g., G. calycosa). In Pheidonocarpa the opposite leaves appear to be joined across the stem by an interpetiolar ridge that develops during matura- tion of the stem. Howard (1 959, 1962) considered the vascular anatomy of the petiole to be important as a taxo- nomic character in many groups. The vascular I 5cm I FIGURE 3.-Leaf bases from one plant of Rhytidopkyllurrr auriculatum: a, petiolatc leaf lacking ?pseudo-stipules?; b, auriculate leaf with decurrent basc; c, auriculate leaf with decurrent base on one side of petiolc only; d, petiolate leaf with ?pseudo-stipules.? trace in the petioles of Gesneria and Rhytido- phyllum has been shown to be a ring or a nearly closed U in cross section (Wiehler, 1970). Wiehler concluded that specific differences may be found if all species were investigated. Attempts to dupli- cate his results showed that the tribe has a distinc- tive petiolar trace pattern; however, the structure of the trace may vary somewhat among the leaves of a single plant depending on age or arrangement on the plant and may not be completely reliable as a specific character. Of the surface characters of leaf blades, the most obvious are the texture and the pubescence. In Q A FIGURE 4.-Trichomcs of Rhytidophyllum: a,b, glandular, R. auriculatum; c, glandular, R. tomentosum; d , nonglandular, R. tomentosum. NUMBER 29 11 Gesneria the adaxial surface is plane, smooth or scabrous, glossy or dull with a thin or thick cuticle, occasionally with large bullae and scattered tri- chomes. Leaves of R h y t i d o p h y l l u m are areolate; each areole bears a single trichome or a group of trichomes, except in R. berteroanum in which the areolae are usually glabrescent. T h e trichomes on the leaves, similar to those found on the stems and inflorescences, are of two types: glandular and nonglandular. T h e glandular trichomes are of three forms. The first (Figure 4a) is short-stalked and at the apex bears cells that have divided anticlinally to the stalk cell into two or more cells (?facherformig,? Sachs, 1915). These glandular hairs probably produce the abundant resinous secretions at the apices of stems and branches. The second type of glandular trichome is a uniseriate hair of two or more cells with a spherical yellow gland at thc apex (Figure 4 b ) , found occasionally along the leaf midveins or on the corolla of many species usually in a ciliate row at the margins of the limbs or scattered at the mouth of the tube, Glandular hairs of a third type were illustrated by Wiehler (1970). These tri- chomes produce mucilage, giving the sticky char- acter to the leaf and stem surfaces in R h y t i d o - phyllum. T h e glands are in a column of two or more cells, with a pointed thick-walled apical cell (Figure 4d) . Nonglandular trichomes are similar in form to the third type of glandular trichome but arise from a basal ring of large epidermal cells (Figure 4c). T h e basal cells often contain deposits of calcium carbonate or calcium silicate (Sachs, 1915), the former detected by evolution of gas bubbles when a weak solution of hydrochloric acid is applied to the leaf. The mineral-impregnated epidermal cells remain after the trichome is lost and give the surface a scabrous or asperate texture as in Gesneria shaferi subsp. depressa. The stalk cells may continue to divide during ontogeny and produce up to thirty cells in a uniseriate column. These usually colorless cells appear white and matted, giving an arachnoid appearance to the surface of leaves of R h y t i d o p h y l l u m acunae, R . lanatum, R. leucomallon, and R. tomentosum. The matted trichomes may decrease water loss (Cutter, 1969). All types of trichomes appear to contain chloro- plasts, at least when immature. The presence of chloroplasts, except in guard cells, is an unusual character for epidermal cells (Esau, 1960). Tri- chomes often contain anthocyanin as well, giving a reddish or purplish color as in some plants of R h y t i d o p h y l l u m leucomallon. Another useful feature of surface morphology is the undulation of the anticlinal walls of epidermal cells. Since the undulation is apparently a geneti- cally determined trait, its characters may have taxonomic value (Linsbauer, 1930). Wiehler (1970) pointed out that the absence or presence and degree of sinuation follows closely the tradi- tional generic lines between Gesneria and Rhyticl- ophyllum. He reports that sixteen species of Gesneria show no sinuation of the abaxial cell walls, four show shallow undulation. All species of R h y t i d o p h y l l u m have deep undulations of the cell wall, except for four species which have no sinua- tion. Examination of some species not available to Wiehler gave results that compare favorably with his results (examples of nonundulating epidermal cell walls can be seen in Figure 5). Stomata, found only on the abaxial surface of leaves, are of the cruciferous or anisocytic type (Esau, 1965) with three or four epidermal cells subsidiary to the guard cells. Arrangement of stomata in groups has been discussed by Sachs (1915), illustrated by Wiehler (1970), and found to vary according to species. Some species not available to Wiehler are illustrated in Figure 5. Gesneria harrisii (Figure 5b) and G . humi l i s (Figure 5f) show tendencies toward grouping of stomata; the other illustrated species appear to have stomata well spaced. Stomata in Gesneria are usually on a level with surrounding epidermal cells, but those in R h y t i d - o p h y l l u m and Pheidonocarpa are elevated to the apex of substomatal domes (Figure 6) outside the spongy parenchyma. Among the examined species of Gesneria, only G . acaulis, G . christii, G. pedi- cellaris, and G. pedunculosa show slight develop- ments of the stornatal domes; these species also tend to have bullate leaves. There may also be a correlation between the development of the stoma- tal domes and the areolate abaxial surface in R h y tidophyllum. The correlation between the presence of the stornatal domes and the tomentose surface of the leaves of R h y t i d o p h y l l u m was noted by Wiehler (1970). Haberlandt (1914) suggests that raised 12 SMITHSONIAN CONTRIBUTIONS TO BOTANY I 1 mm I FIGURE 5.-Abaxial leaf epidermis in species of Gesneria: a , G . clandestina; b, G. harrisii; c, G. ventricosa (from Dominica); d, G. haitiensis; e, G. fruticosa; f, G. humilis (from Haiti). NUMBER 29 13 stomata usually occur in plants growing in shady habitats, and because the stomata are elevated, transpiration probably increases due to a freer flow of air over and around the stomata. Biebl and Germ (1950) also suggest this reason for the presence of stomatal domes in plants in humid environments, Since R h y t i d o p h y l l u m grows in dry exposed areas on banks (Figure 1 b ) or in clearings, this reasoning is unlikely. I t is likely, though, that because of the dense trichome covering of these plants, the microclimate at the leaf surface is hu- mid, and the stomata, raised on stomatal domes, may allow more efficient transpiration in the microclimate. The internal anatomy of the laminae of mem- bers of this tribe has not been studied since the investigations published in 1915 by Sachs. An attempt has been made to examine some of the species she studied as well as others not available to her. The adaxial epidermis appears one-cell thick in investigated species of R h y t i d o p h y l l u m auricu- l a t u m , R. berteroanum, R . leucomallon, R. tomentosum, Gesneria acaulis, G . christii, G. citrina, G. libanensis, and Pheidonocarpa corym- bosa. Gesneria pedicellaris, however, a close relative of G. acaulis and G. christii, has some leaves with a uniseriate row of cells and others with two or more layers in an apparent multiple epidermis. Although detailed developmental studies have not been done to determine the true identity of the subepidermal tissues, layers of cells resembling a multiple epidermis, which may decrease water loss from the mesophyll (Cutter, 1969), are distinctive features observed in cross sections of leaves of I O.5mm I FIGURE 6.--Stomatal domes on thc abaxial leaf surface of Rhytidophyllum tomentosum. many species of Gesneria. This adaxial tissue of large thin-walled water-storage cells was termed a hypodermis by Sachs (1915), but because of its dissimilarity to the mesophyll below and similarity to the epidermal layer and occasional alignment in radial rows, which may reveal the origin of the cells by periclinal divisions from the epidermis, the layers of multiple epidermis appear to have been derived from the protoderm. According to Esau (1965) the periclinal divisions that produce multiple epidermis may occur at different stages of leaf growth but usually when a leaf is several inter- nodes below the apex. Thus, leaves taken in dif- ferent stages of maturity may differ in amount of the subepidermal tissue. Light intensity may determine the relative size and number of rows of cells or layers in the meso- phyll. Gesneria exserta, which grows in nearly full sun, has a thick palisade layer of one to three rows of elongated cells. The lower layers are more loosely joined and become more isodiametric in apparent transition to the spongy parenchyma. Spongy mesophyll cells near the lower epidermis have lateral extensions allowing contact between the cells and spaces which may allow gas exchange. Gesneria reticulata, a species that does poorly in full sun, has a single layer of palisade parenchyma and only one or two layers of spongy mesophyll. In this latter species, however, beneath the spongy cells is a region of abaxial multiple epidermal cells, differing from the spongy mesophyll cells in being larger and lacking chloroplasts, but like them in being bounded by air spaces. Scattered among the cells of spongy mesophyll in Gesneria pedicellaris, at least, are crystals or mineral deposits as large as the cells (this observa- tion is in contrast to Sachs? statement that large crystals do not occur in this family). These differ in shape and size from the usual calcium oxalate druses and may be calcium carbonate or calcium oxalate as found in trichomes. Other cells, par- ticularly near the veins, contain raphides similar to those found in the pith of this species (and reported from other species by Sachs). The minor veins in the leaf are composed of a few xylem elements subtended by phloem, all surrounded by a sclerenchymatous bundle sheath. In Gesneria pulverulenta the bundle sheath pro- jects like a pillar to the base of the adaxial multi- ple epidermis. In R h y t i d o p h y l l u m the minor 14 SMITHSONIAN CONTRIBUTIONS T O BOTANY veins have a cortical sheath causing a prominent bulge in the leaf surface. The vascular pattern of the midveins is similar to the patterns in petioles. Main veins of most species are U-shaped rows of xylem vessels with an abaxial layer of phloem. Within the U-shaped strand are scattered arcs or bundles of xylem vessels and abaxial phloem layers. Exceptions are found in Gesneria viridiflora subsp. quisqueyana, which has three concentric U-shaped strands with intervening layers of parenchyma, in G. pedicel- laris with a nearly closed cylinder, and in R h y t i d o p h y l l u m species where the vascular trace was shaped like an inverted capital omega: a. The cortex surrounding the vascular strands is composed of parenchyma, with scattered sclerotized cells revealing large pit-pairs. Sclerotized cells also form a large area at the apex of the main veins, -* I- I I X I 0 W Y 0 w Y FIGURE 7.-a, Schematic drawing of inflorescence; b, inflores- cence diagram. (Terminal flower = solid circle: lateral flowers = open circles; bracts = curved lines; internode be- tween bracts at base of terminal flower and bracts at apex of primary peduncle = X.) and form a one- to three-celled ring around the outer margin of the vein. Layers of thick-walled cells occurred on the abaxial side of the phloem in Rhytadophyl lurn auriculatum. Iwa-oR~scmcEs.-Inflorescences in the tribe Ges- nerieae are basically similar; those in each species can be shown to be examples of a stage in a reduction series from a compound dichasium to a single flower. The simplest inflorescence is a single flower borne on an axi5 with two lateral bracts (Figure 15), all arising from a leaf axil. The most complex inflorescence is that shown in Figure 7, a com- pound dichasium with a terminal flower on a central axis, which bears two sets of bracts: one set at the apex of the primary peduncle in whose axils are borne lateral axes, which simulate the central axis; the other set of bracts are at the base of the terminal flower. One of the latter bracts is fertile and bears a branch system of flowers similar to a w n n n b - 0 0 (0) (0) ((0x0) (0x0)) (0) (0) 0 0 v Y v v 0 - FIGURE &--a, Schematic drawing of inflorescence; 6, inflorescence diagram. NUMBER 29 15 that of the lower lateral branch axes. The other bract is apparently sterile, but in the putative natural hybrid between Gesneria acaulis and R h y t i d o p h y l l u m t o m e n t o s u m a flower is produced whose pedicel is adnate to the pedicel of the terminal flower for a distance u p to 6 mm; how- ever, this adnate flower is placed between the terminal flower and the branch system rather than on the side opposite the branch. One or both bracts at the base of the terminal flower may be lacking, or one may be adnate to the branch. The branch at the base of the terminal flower may be reduced to a simple dichasium. The internode between the bracts at the base of the terminal flower and the bracts at the apex of the primary peduncle (labeled ?X? in Figure 7) is usually reduced so that a terminal flower, its bracts and branch system, as well as the lower branches appear to arise from the same point. This internode re- duction is common and persistent in all stages of the series. T h e inflorescence just described and illustrated in Figure 7 is found in nearly all species of R h y t i d o p h y l l u m but is known only in a variety of Gesneria, G. scabra var. sphaerocarpa. A slight reduction is found in R h y t i d o p h y l l u m onacaense (Figure 8). The bracts of the terminal flowers of the main and lateral axes are occasion- ally fertile, but bear single flowers and never branch systems at that point. Abortion of the bracts at many points is common (their presence is FIGURE 9.-u, Schematic drawing of inflorescence; b, inflorescence diagram. sometimes seen as vestigial bumps or swellings), but always a pair is present at the base of the lowest lateral axes. From the next stage of the reduction series to the final single-flowered inflorescence, the inter- node ?X? is lacking, as is one lateral flower in the terminal dichasium. The remaining two flowers are decussate to the lateral branch axes. The order of flowering follows the pattern one might expect in a dichasial system: the terminal flower of the central axis opens first, followed by the remaining flowers in the terminal set of bracts, then the terminal flowers of the secondary branch axes and their subsidiary flowers, then the four terminal flowers of the tertiary axes, and so on to the limit of the development of the inflorescence. In many b 0 FIGURE IO.-u, Schematic drawing of inflorescence; b, inflorescence diagram. 16 SMITHSONIAN CONTRIBUTIOKS T O BOTANY species, particularly Rhyt idophyl lurn berteroanum, the inflorescence ceases further growth and devel- opment when a flower is fertilized. Gesneria fruticosa (Figure 9) has four bracts at the base of the terminal flower and its subsidiary flower where it joins the lateral axcs. The number of bracts is reduced to two in Figure 10, the struc- ture is found in many species, including Gesneria acaulis, G . humil is , and G. viridiflora subsp. q u isq uey ana . By reduction of the lateral branch axes in Fig- ure 9 to single flowers and maintenance of the four bracts, the inflorescence found in Gesneria fxd?cellaris (Figure 11) can be derived, a structure of two pairs of decussate bracts with flowers in their axils. The inflorescence in Figure 12 could result either by reduction of lateral branch axes shown b 0 - FIGURE 11 .-u, Schematic drawing of inflorescence; b, inflorcscence diagram. in Figure 9 to single flowers or by reduction in the number of bracts from the inflorescence shown in Figure 11. GeJneria ventricosa displays this type of inflorescence in the typical subspecies. Further reduction in number of flowers is obvious in Gesneria pedunculosa (Figure 13) or G. purnila subsp. mirnuloides. The usual number of flowers is two: the terminal flower and its sub- sidiary. The lateral flowers are seldom present though the bracts pcrsist. Reduction to a single flower is through loss of the subsidiary flower. I n Gesneria cuneifolia (Fig- ure 14) the reduction probably parallels and sur- passes Gesneria pedicellaris (Figure 11). The lateral and subsidiary flowers have aborted, leaving the terminal flower with four bracts at its base. In Gesneria calycosa the reduction to a single flower presumably occurs after the abortion of the bracts at the base of the terminal flower (Figure 15). The bracts that remain on thc flower stalk 0 a b (0 0) 0 FIGURE 12.-u, Schematic drawing of inflorescence; b, inflorescence diagram. NUMBER 29 17 probably subtended the lost lateral axes. This last inflorescence example may be considered most ad- vanced in the genus, for i t is usually found in those species which have the, presumably, more-recently evolved bat-pollination mechanism. FLOWERS.-AII species in this tribe have flowers similar in structure-i.e., a floral tube in the sense of Wilson (1974b) composed of the adnate bases of the calyx, corolla, and stamens fused into a cup surrounding and adnate to the inferior or half- inferior 2-carpellate unilocular ovary. Above the attachment to ovary, the flower parts making up the floral tube become distinct. The calyx has five lobes sometimes connate briefly at the base. The five petals of the corolla are fused into a tube with a five-lobed limb at the apex. Adnate, at least briefly, to the base of the corolla tube are 'four stamens and a staminode. The free calyx lobes may be sulcate at base and awned above (e.g., Gesneria onychocalyx), flat- tened as in G. pedunculosa, or terete (e.g., G. duchartreoides), glabrous or pubescent, but usually green. One to three veins in each persistent lobe may become prominent in the fruit. The calyx occasionally surpasses the corolla, as in G. calycosa, or may be reduced to less than 1 mm long in G. cubensis var. truncata. In those species where resin is abundant on the young leaves, the calyx is also very resinous. The shape of the corolla tube may be an adap- tation to the pollination mechanism-i.e., large and campanulate in suspected bat-pollinated species (Figures Ih-i, 16b-c,m, 22), or narrow and tubu- lar in the hummingbird-visited species (Figures 16d-k, 21a). One collection of Rhytidophyllum auriculatum (G-1066) bore at the same time corollas of both shapes, exhibiting a peloric trait found also in some species of Columnea, Rech- steineria, and Sinningia (H. E. Moore, pers. comm.). Green corollas, due to chloroplasts in the 0 FIGURE 13.-a, Schematic drawing of inflorescence; b, inflorescence diagram. 0 b FIGURE 14.-a, Schematic drawing of inflorcsccnce; b, inflorescence diagram. 18 SMITHSONIAN CONTRIBUTIONS T O BOTANY mesophyll cells, are common and probably are due to nondevelopment of anthocyanins. Protogyny and protandry both occur among the species of the Gesnerieae. The dehiscence of the anthers and the receptivity of the stigma are usually separated by a day, but environmental conditions may cause an overlap, allowing the possibility of self-pollination. The adnation of the filaments to the base of the corolla has been used by many authors as the main distinguishing character between Gesneria and R h y t i d o p h y l l u m . Unfortunately, the trait is not always definite, but seen in combination with characters of the fruit, habit, and foliage, filament adnation is still a valuable marker. Filaments are nearly straight in the tubular species and curved or geniculate in the species with campanulate corollas. Often the base of the filament is invested with pilose colorless trichomes. Filaments and anthers may be white, green, yellow, or reddish depending on the species. The filament or con- FIGURE 15.--a, Schematic drawing of infloresccnce: b, inflorescence diagram. nective may be prolonged in some species beyond the anther; the connective appears to be quite variable and may not be as diagnostic here as it is in some species of Codonanthe (Moore, 1973a). Anthers are usually coherent in two pairs by their apices, arranged in a row of four side by side, or separate. The coherence is tenuous in some species and anthers may become free after anthesis. Dehiscence of the anthers is longitudinal, and occurs soon after anthesis in protandrous species (e.g., Gesneria pedunczilosa), or about a day later in the protogynous species (e.g., G. viridiflora subsp. quisqueyana) . Exsertion of the stamens seems to be a detail of the pollination mechanism and may effect a reproductive barrier that may prevent cross- pollination among sympatric species. There ap- pears to be evidence of coevolution and adapta- tion in some species (see Pollination and Seed Dispersal). In Gesneria section Uuchartrea, sus- pected as being bat pollinated, thc stamens are included, e. g., Gesneria viridiflora subsp. quis- queyana (Figure I 6 m ) ; if a bat probed the flower for nectar, pollen would be deposited on the fore- head of the animal. In some species of Gesneria section Dittanthera, where the stamens are long- exserted, e.g., G. pedunczilosa (Figure li) or G. calycosa (Figure 22a), a probing bat would receive a coating of pollen at the back of the head. In the plants believed to be pollinated by hum- mingbirds, a similar state exists: flowers of Ges- neria section Gesneria have included stamens from which pollen may be shaken or brushed onto the top of the base of the bill of a hummingbird visitor; Gesneria section Pentarhaphia includes plants with exserted stamens from which pollen may be shaken or brushed onto the top of a bird?s head (Figure 1 6 4 . The styles of all these plants appear to be approximately the same length as the stamens and terminate in stigmas so oriented that pollen from an animal?s head may adhere to them. Early pollen studies in the Gesneriaceae were executed using light microscopy and were devoted either to development (Goldmann, 1848) or to casual observations of surface features (Schnizlein, 1843-1870). The only critical study of the pollen morphology of the Gesneriaceae since a brief sur- vey in 1952 by Erdtman has been the unpublished investigation by Pauline Woods at the Royal NUMBER 29 19 Botanic Garden, Edinburgh. To determine if a closer look would reveal additional information, I examined 27 examples of pollen from the tribe Gesnerieae with a scanning electron microscope. Pollen among the species of the tribe Ges- nerieae is relatively uniform as seen under the scanning electron microscope. Although surface topography did display some variation among species, obvious differences in size and shape were not seen; the size limits in this group are well within the limits of the family as observed by Erdtman (1 952). Pollen grains are isopolar with a nearly circu- lar amb, tricolpate with three mesocolpia and two truncate or rounded apocolpia, prolate; size is small to medium (18.5-27.0 pm long at the polar axis, 8.5-6.0 pm wide at the equatorial axis); sexine is usually reticulate with lumina heterobro- chate or rarely homobrochate, about 1 pm across, muri about 1 pm wide, seldom nearly smooth or verrucose; colpi are long and narrow. The exceptions to the typically reticulate sexine surface among the 27 examples investigated were found in Gesneria scabra var. sphaerocarpa, with a sexine that appeared to be nonreticulate and nearly smooth (Figure 17f), and in Gesneria clandestina (Figure 18a) and G. fruticosa, both of which had a verrucose or warty sexine. Gesneria uiridifiora subsp. sintenisii (Figure 17a) had a rugose outer surface that appeared intermediate between the verrucose type and the typical reticu- late forms. The latter three species have apparently become modified morphologically in corolla char- acters related to bat pollination. Perhaps the verrucose or warty surface is a modification in the pollen associated with the more recent bat pol- FIGURE 16.-Flowers of Gesneria and Rhytidophyllum: a , artificial intergeneric hybrid Rhytido- phyllum leucomallon X Gesneria pedunculosa, G-1232; b, R . auriculatum, G-905; c, R . leucomal- ion, G-1010; d , Gesneria ventricosa subsp. ventricosa, G-940; e, G. christii, G-1008; j , G. pedicel- lark, G-898; g, G. acaulis var. glabrata, G-876; h, G. cuneifolia, G-869; i, G . citrina, G-888; j , G. pulverulenta G-1034; h, G. paucipora, (3.769; 1, G. humilis, G-1365; m, G. viridiflora subsp. quisqueyana, G-903. 20 SMITHSONIAN CONTRIBUTIONS T O BOTANY FXCURE I?l.-Pollen grains of tribe Gesnerieae as seen with scanning electron microscope, scale indicated by ruled line: a, Gesneria viridiflora subsp. sintenisii (Talpey 23) , scale = 5 pm; b, G. Pedicellaris (G-898), scale = 5 pm; c, G. acaulis var. glabrata (G-876), scalc = 10 pm: d , G. acaulis var. acaulis (G-877), scale = 10 km; e, Xhytidophyllum tomentosum (G-841), scale = 10 pm; f, Gesneria scabra var. sphaerocarpa (G-881), scale = 10 pm; g, G. uentricosa subsp. ventficosa (G-940), scale = 10 pm: h, G. humilis (Alain Liogier 4434) , scale = 5 bm. NUMBER 29 21 FIGURE 18.-Pollen grains of tribe Gesnerieae as seen with scanning electron microscope, scale indicated by ruled line: a, Gesneria clandestina (Harris & Brit ton 10679), scale = 10 pm; b, G. viridifiora subsp. viridipora (Clemente 5089), scale = 2 pm; c, G. pulverulenta (G-1034), scale = 20 pm; d, G. cuneifolia (G-763), scale = 10 pm; e, G. paucifiora (G-769), scale = 10 pm; f, G. humilis (G-1365), scale = 3 pm; g, putative natural hybrid Gesneria acaulis X Rhytidophyl- l u m tomentosum (G-1371), scale = 10 km; h, Pheidonocarpa corymbosa subsp. corymbosa (G-1367), scale = 10 pm; i, Gesneria pedunculosa (G-767), scale = 10 pm; j , G. calycosa (G-1037), scale = 10 pm; k, G. pumi la subsp. pumila (G-874), scale = 10 pm. 22 SMITHSONIAN CONTRIBUTIONS T O BOTANY lination; warts may allow the grains to adhere to the fur of the flying mammal. Other species probably modified for bat visitation, Rhytido- phyllum tomentosum (Figure 17e), Gesneria calycosa (Figure ISj), and G. viridiflora subsp. viridiflora (Figure 18b), have not developed this surface topography, but have grains with the reticulate surface typical for the tribe, The startling appearance of the pollen of hybrid forms added evidence of their origin. Pollen of two natural intergeneric hybrids, Rhytiddphyllum to- mentosum X Gesneria scabra var. sphaerocarpa and Rhytidophyllum tomentosum X Gesneria acaulis var. glabrata (Figure 18g), showed distor- tion and general collapse of the grains. A hybrid between two species of Gesneria, G. pedicellaris X G. pedunculosa, did not show this collapse. Pollen of Gesneria humilis from Haiti also showed collapse and distortion similar to I the hybrids (Figure I8f); however, pollen from the same species from Cuba appeared to be normal (Figure 17h). The remainder of the pollen ex- amined appeared to be relatively uniform in struc- ture but differed somewhat in size. The half-inferior to inferior bicarpellate, uni- locular ovary is hemiparacarpous according to Ivanina (1965). According to Wilson (1974a) the two carpels are arranged as in the closely related Scrophulariaceae and ?lie in an anterior-posterior position with relation to the floral axis and not, as stated by Hutchinson (1959), to the right and left of the floral axis.? Earlier authors (e.g., Lawrence, 1951) have written that the placentation in the Gesneriaceae is parietal with placentas bearing numerous anatropous ovules. Weber (1 97 l), how- ever, has shown that in Gesneria libanensis the ovules are borne only on the surfaces of the in- truded placental lamellae and not on the outside walls of the carpels. In Pheidonocarpa corymbosa the apex of the half-inferior ovary protrudes be- yond the disc and becomes accrescent to form a beak in the fruit, an obvious character that sep- arates this species from Gesneria. The disc of all species is apparently a ring of fused glands, be- coming five-lobed, sinuate, or unlobed. According to Brown (1938) the disc in the Gesneriaceae is nectarial. In plants like those in the tribe Ges- nerieae, Brown suggests that the inferior ovary, but not the nectariferous tissue, has been sur- rounded by the torus or receptable; Moore and Lee (1968) suggest the possibility that the disc in some genera may be a reduced inner whorl of stamens and call attention to the need for de- tailed study. However, the disc in Gesneria bears glandular trichomes that exude abundant nectar in many species and may testify to its nectarial origin. Wilson (1974a, 1974b), after extensive study of nectarial tissues in the Gesneriaceae, con- cluded that the disc ?is an enation; there is no evidence that it is a modification of any pre- existing cycle of floral parts.? In Gesneria pedunculosa, a protandrous species, the style is patent during the first day after an- thesis, but becomes erect during the second day. In protogynous species, e.g., Gesneria viridiflora subsp. quisqueyana, the style is exserted while the corolla is opening and before the anthers dehisce. The papillate stigma is capitate and enlarged in Gesneria humilis, slightly enlarged in most species of Rhytidophyllum, stomatomorphic in most spe- cies of Gesneria, but in Pheidonocarpa the stigma is prominently bilobed. FRvIT.-capsules are elongated and h e a r in Gesneria calycina, turbinate in most other species of Gesneria, or nearly spherical as in most species of Rhytidophyllum. The rostrate apex appears only in Pheidonocarpa. Dehiscence occurs usually through the median radial splitting of the carpels into two or four valves from the apex. An unusual fruit referred to here as a splash cup develops in some species of Gesneria (see Pollination and Seed Dispersal). In Gesneria citrina, G. cuneifolia, G. humilis, and others the apex of the capsule splits or withers to form a round opening and exposes seeds, which may be ejected by rain or waterfall splash. In many species of Rhytidophyllum, the capsule is inverted during maturation allowing seeds to fall out when the fruit opens. The veins of the floral tube surrounding the ovary in species of Gesneria usually cnlarge to form prominent costae on the capsules; Pheidono- carpa and Rhytidophyllum species do not show this character. SEEDs.-The seeds develop from anatropous unitegmic ovules (Davis, 1966). Seeds of all speci- mens investigated were similar in size, averaging about 800 pm long and 250 pm wide, but ranging from 1300 pm long in Gesneria calycosa (Figure 199) to 500 pm long in G. pumila subsp. neglecta, from NUMBER 29 23 FIGURE 19.-Seeds of Gesnerieae as seen under a light microscope: a, Gesneria heterochroa (Ekman 8741); b, G. acaulis var. acaulis (G-877); c, G. christii (G-1008); d, Rhytidophyl- lum grande (Proctor 18312); e, Gesneria acaulis var. glabrata (G-876); j , G. calycina (Proctor 22125); g, G. calycosa (Proctor 21517); h, Rhytidophyllum tomentosum (G-841); i, Gesneria ventricosa subsp. ventricosa (C-940); j , G. humilis (G-1365); k , G. cuneifolia (G-857); I , Rhytido- phyllum grande (Proctor 16645); m, Gesneria alpina (Proctor 9616); n, G. duchartreoides (Alain Liogier & Lopez F . 7318). 24 SMITHSONIAN CONTRIBUT LONS T O B O T A S Y FIGURE PO.-Seeds of Gesnerieae as seen with scanning electron microscope (scale = 100 pm): a, Gesneria christii (G-1008); b, G. humilis (G-1365); c, Rhytidophyllun tornentosurn (G-841); d , Gesneria cuneifolia (G-857). 450 pm wide in G. acaulis (Figure 19b,e) to 150 pm wide in G. scabra var. fawcettii. Seeds are spindle-shaped (Figure 1 Sa,d,h,i,k-n), linear (Figure 19f-g), rhombic (Figures 19i, ~ O C ) , or oblong (Figures 19b-c, 20d). All appear twisted or bear spiral lines, which are apparently the remnants of the cell walls of the integument. At maturity the seeds probably twist upon drying. The walls of the outer cells appear inflated in species of Gesneria (Figure 20a-b,d) but sunken and less twisted in Rhyt idophyl lurn (Figure 20c). Ges- neria ventricosa, however, appears intermediate in this character. Upon germination, the seedlings are isocotylous, consistent with the division by Burtt (1962) of the Gesneriaceae into two subfamilies based on seed- ling characters. CONCLUSIONS T h e main purpose for examining the species of the tribe Gesnerieae was to determine whether thcre are differences that could distinguish the genera Gesneria and R h y t i d o p h y l l u m consistently. Plants in these genera have been considered con- generic by many authors, e.g., Linneaus (1753), Baillon (1888), Kuntze (1891), and more recently a union was proposed by Wiehler (1970). In recent floras of the West Indian region (Adams, 1972; Morton, 1957b), these genera have been separated by thc length of the adnation of the filaments to the corolla tube, a distinction noted by Fritsch (1894). Other characters may now be added. Table 1 gives the characters that distinguish Gesneria, R h y t i d o p h y l l u m , and Pheidonocarpa. Consideration of these characters and observations of the plants in tlic field have led to the conclusion that Gesneria and R h y t i d o p h y l l u m should be maintained as separate genera. Thcre is overlap in many characters, and no single character may distinguish all taxa in one genus from all taxa in SUMBER 29 T A B L E 1 .-Comparison of characters of genera in the Gesnerieae 25 Charm ter Habit Habitat Crystals LeaYeS arrangement "pseudo-stipules" adaxial surface glandular muci- laginous hairs "on-glandular uniseriate hairs multinle epidermis abaxial epidermal cell wall undulation minor veins stornatal domes stornatal 'islands' Inflorescence Flowers filament adnation ovary capsule shape costae seed walls GeS"erI.3 Rhytidophyllum Pheidonocarpa acaulescent herbs shrubs to trees subshrubs subshrubs to trees ~ usually protected =aphidesa a1 terns t eb absent plane, smooth & nitid, scabrous, O C C ~ S . bullate absent few-celled most species none or shallow no cortical sheath seldomd many species reduction series (Figures 7-15) less than thickness of filament base inferior linear, turbinate rarely spherical present, if not, then capsules "arruco8e usually inflatedf exposed marginal styloid alternate occasional areo1ate present up to 30 cells arachnoid surface absent usually deep' cortical sheath ?.lw.%ys few speciese compound dichasium (Figures 7-8) more than thickness of filament base inferior usually spherical absent sunken exposed limestone cliffs styloid opposite, decussate absent scabrous absent few-celled absent none unknown always all species compound dichasium (Figure 8) more than thickness of filament base half-inferior turbinate, long curved beak absent or obscure sunken a Gesneria pauciflora has styloid crystals. Leaves of Gesneria fruticosa are opposite and decussate in one population. NO cell wall undulation in Rhytidophyllum earlei, R. lomense, and E. rupincola. Slight stornatal dome development in a few species of Gesneria. Stomata1 'islands' present in Rhytidophyllum - and g. rupincola. Gesneria ventricosa is trangitional. -~ the other. Evidence of their distinctness may be found also in pollen characters of intergeneric hybrids: collapse of pollen grains observed in scanning electron microscopy, and the low pollen stainability, compared to noncollapse of pollen grains in an interspecific hybrid in Gesneria, which has high pollen stainability. Gesneria corymbosa and G. mortonii should be removed from Gesneria, based on field observa- tions and laboratory investigation, also sum- marized in Table 1. Three solutions are suggested for the disposition of the two species: ( 1 ) reunion with Heppiella, disqualified for the reasons de- tailed by Wiehler (1971); (2) transfer to Rhytido- phyl lum with which many of the characters appear to correlate more closely than with Gesneria, but the species would still remain an anomalous group within that genus, and this solution is therefore rejected; (3) erection of a new genus in the tribe Gesnerieae. T h e chromosome number, structures of inflorescence and flowers, and leaf anatomy all point to a closer relationship to Gesneria and Rhytidophyllurn than to other genera. This action, however, would destroy the traditional tribal 26 SMITHSONIAN CONTRIBUTIONS T O BOTANY character of a1 ternate leaf arrangement. Emenda- tion of the tribal circumscription is therefore necessary to account for the addition of the new genus, Pheidonocarpa. This third course of action is taken herein. Pollination and Seed Dispersal The evolution of flowering plants has probably been largely the result of adaptive pressure toward efficient methods of dissemination of pollen and seeds. This section deals with these aspects of dis- semination in the Gesnerieae, primarily in the genus Gesneria; (1) the pollination biology and probable coevolution 01 Gesneria species and vertebrates, and ( 2 ) seed dispersal and its primary reliance on water and winds. MATERIALS AND METHODS T o correlate the sizes and shapes of flowers to possible hummingbird and bat pollinators, meas- urements of suspected animal visitors were taken from specimens in the collections of Cornell University, and in the Divisions of Birds and Mammals at the National Museum of Natural History, Smithsonian Institution, Washington, D. C. TABLE 2.-Measurements of hummingbirds Species 2.4 2 . F 2.i 2.0 1.1 1.5 1 . 7 1.9 ; .3 I , i 3.0 L . 2 1.4 2.7 2.1 2.8 2 . 7 L.2 2.0 1.2 1.8 2.0 2.3 222 3.0 5 . 3 2 . 0 1.8 2 . 8 3 . 4 For hummingbirds, the following data were collected: (1) the name of the species, ( 2 ) the collection location, (3) the accession number, (4) the length of the bill measured in cm, and (5) the width of the bill (in mm) at the base. Averages of these measurements are shown in Table 2. Selected specimens were photographed for a record. Because of the varied shape of heads among the species of bats, more extensive data were collected: (1) the species name, (2) the collection location, (3) the accession number, (1) dorsal-ventral width in mm of head at middle of eye, (5) length along dorsal rostrum from level of middle of eye to nose leaf, (6) depth in mm of nose from base of nose leaf to bottom of lower jaws, (7) width in mm across skull at center of eyes, (8) width in mm across anterior tip of nose, (9) depth in mm ver- tically from crown of head to lower jaws, and (10) distance in mm from tip of nose leaf to bottom of lower jaws. For comparison with corolla length and width, it was found that meas- urements 4 and 5 were critical. Averages of all of these measurements are shown in Table 3. Again, selectcd specimens were photographed. POLLINATION BIOLOGY By far the greatest part of the literature on co- evolution of plants and animals has been devoted to the field of study known variously as pollination biology, pollination ecology, or anthecology. This field of study began over 200 years ago with the work of Kolreuter (1761), was expanded by Sprengel (1793), Muller (1883), and Knuth (1906-1909), but only in the last few decades has yielded information to show the effects of adapta- tion and selection resulting in speciation. T h e works by Faegri and van der Pijl (1966, 1971) summarized the recent work in pollination biology. Current studies of pollination biology rest on a concept of coevolution or development of mutual- istic relationships between flowering plants and animals. According to Baker and Hurd (1968), coevolution or reciprocal evolution among plants and animals occurs in a series of steps, a result of natural selection toward the most efficient model for reproduction in both partners. An animal that can derive food or energy from a plant with a smaller expenditure of energy than another animal NUMBER 29 27 is more likely to survive to pass on its hereditary characters. Likewise, a plant that is more attractive to potential pollinators or seed dispersers is more likely to have its hereditary characters appear in the next generation. Because of their ?tremendous evolutionary plasticity? and coevolution with animals, ?angiosperms have attained the highest level of organization in the plant kingdom? (Takh taj an, 1969). Since pollination by abiotic means, such as wind or water, is neither known nor suspected in Ges- neria, only biotic pollination will be discussed. Three methods of biotic pollination are suggested and will be considered: transfer of pollen via insects, birds, or bats. Self-pollination may be im- portant in some species and will be mentioned in relation to distribution patterns. Flowers may have a combination of characters to attract more than one pollinator, as well as a capacity for self-pollination. Baker, Cruden & Baker (1971 :1127) recently wrote: ?very few plants have only one kind of visitor capable of effective pollination . . . and practically none have only one kind of visitor capable of gaining sustenance from the floral parts or the nectar.? Thus, while flowers of many species of Gesneria may be pollinated by bats, these same flowers may be visited by hum- mingbirds or insects. Figure 21b is a photograph of a hummingbird (Anthracothorax dominicus) visiting Rhytidophyllum auriculatum, a species probably adapted for bat pollination. Among the invertebrates, only insects may be considered as possible visitors to Gesneria flowers. The interactions of many plants and insects have been thoroughly studied: extensive observations are detailed in the early works by zvvcv (1883) and Knuth (1906-1909), and a summary discussion of coevolution of plants and insects has recently been published by Macior (1971) along with an extensive bibliography on the subject. Insect pollinators are usually members of the orders of Coleoptera, Diptera, Hymenoptera, and Lepidoptera. The characters of the flowers at- tracted by insects have been listed by Faegri & van der Pijl (1966, 1971). Flowers specialized for visitation by one insect taxon are likely to be less attractive to other groups of insects, i.e., flowers attractive to bees may differ from flowers attractive to flies, butterflies, or moths. There are no previously documented reports of insects visiiing flowers of Gesneria. A comparison of the characteristics of Gesneria flowers with those of insect-pollinated flowers given by Faegri and van der Pijl (1966, 1971) shows that Gesneria is not likely to be pollinated by insects. The genus lacks blue or pure yellow flowers especially attrac- tive to bees, ultra-violet reflecting nectar guides used by bees and flies to locate nectar or pollen, a strong fruity odor attractive to beetles or moths, radial erect flowers with a landing area attractive to butterflies, regular simple flowers visited by flies, and the versatile anthers of moth-pollinated flowers. The tubular or campanulate flowers dis- courage beetles. However, while collecting in the Dominican Republic, I saw honeybees (Apis mellifera) enter flowers of Rhytidophyllum ber- teroanum in the field; the honeybees that visited the flowers of this species could not be the original pollinators, for they are recent immigrants to the New World. The tubular, red flowers of R. berteroanum appear to be adapted to humming- bird pollination. In species of Gesneria with tubu- lar, red corollas, bees may occasionally be attracted to the nectar, which is usually available in abundance. Because the anthers are positioned on the upper side of the corolla tube, as in the campanulate-flowered species, and from observa- tions of Apis mellifera entering the tubular corolla TABLE 3.-Measurements of bats 1.2 1.2 1.0 1.5 0.9 * * F b e r t o RiCo Cuba i . O 1.: 1.1 1.5 0.9 1.6 * 3cminroan Repcb1.c 1.2 1.0 0.3 1.3 0.8 1.8 * P u e r t o H 1 ~ 3 0.8 0.6 0.5 0.8 3 . E 1.2 0.7 :ia.tL 0.3 C . 3 O.b 0.8 0 . 6 1 . 3 0.8 OJba 0.8 0.9 C.6 O . e 0.5 1.4 * Jamdl C B 0.6 0.9 C . E 0.8 0.5 1 . 3 0 . 7 Jnnai c a 0.9 0.8 3 . c 3 . 3 0.6 1 . c 1.0 CLbZ, *&I:; 0 . 8 3 . 3 0 . 5 3 . 3 0.4 1.1 0.6 e l - v e n t r a l wrcth of read m d d l c c f e y e , 2 = l c n g ; 7 z i o n g acrsaL L P l e v e l of n ; d d l c of e y e n o s e leaf; 3 = ? e p t h of nose from base c a c r o s s skrl: a?. Centor C C e y e s ; 5 = r i d t ?7,ert:oal:y from CIOYIT. 0: hew2 TC l s s c r J bot:om of loaer jaas. * ~ n m neasm~red, (X) = prescrbec s p e c i o e n s n o t a v i : l a b l e , neasurener.%s d e r i v e d f r o m S k u l l . 28 SMITHSONIAN CONTRIBUTIONS T O BOTANY of Rhytidoplayllum berteroanum from the lower side, it appears that bees would seldom contact the anthers and stigma to transfer pollen. Therefore, bees are not likely to be important pollinators. If insects are eliminated from the likely princi- pal pollinators of Gesneria, one must search among the vertebrates to find the probable pollinators. According to Allen (1939) and Faegri and van der Pijl (1966, 1971), vertebrate pollinators are pre- dominantly phenomena of the tropics, for only in tropical regions with their evenness of climate are blossoms available for food at all times during the year. In the neotropics, two vertcbrate pol- lination mechanisms appear to predominate: hum- mingbird pollination and the more recently evolved and less well-known pollination by bats. Pollination by birds was first described by FIGURE 21.-Hummingbirds with flowers they may visit and pollinate: a, Anthrucothorux dorninicus from the Dominican Republic with Gesneria pedicellaris (top), G. christii (middle), and G. reticulata (bottom); b, Anthrucothorax doininicus visiting Rhytidophyllum uuriculatum in the Dominican Re- public (arrow indicates pollen at thc base of bill; photograph by T. E. Talpey). Trelease in 1881 and was later studied by Fries (1903) and Werth (1915) among others. The work by Grant and Grant (1968) has elucidated many of the mechanisms in ornithophily. Ornithophilous plants are usually visited and pollinated by birds of the families Coerebidae, Nectariniidae; Meliphagidae, Drepanididae, or Trochilidae (Welty, 1963; Wallace, 1963). In the Trochilidae, the hummingbirds are highly adapted to taking nectar from flowers by their ability to hover, a visual sensitivity to red, a long bill, and a tongue modified for taking up nectar. Characters of bird-pollinated flowers are listed in Table 4. These characters in combination serve also to exclude other possible competitors for pol- len or nectar. The red color of these flowers is seen as the unifying character of hummingbird- pollinated flowers. Baker (1961:66) stated that the red color with its longer wavelengths makes the flowers more easily seen against the darker foliage at dusk or early in the morning. Grant and Grant wrote that the common red color of ornitho- philous flowers is the result of mimicry or adaptive coloration. Hummingbirds appear to associate red color with large amounts of nectar. Since most insects and bats are not attracted to a bright red color (unless combined with ultraviolet) and are thus excluded from pollination, the plants are dependent upon hummingbirds for pollination. The stages in coevolution of birds and flowers cannot be detailed with certainty, but Grant and Grant (1968:87) suggest that a preadapted ?swift- like bird-tropical, insectivorous, and agile on the wing? was the probable ancestor for the nectar- feeding hummingbird. The plants have probably developed from a preadapted insect-pollinated ancestor, but since there are no known present-day exclusively insect-pollinated species of Gesneria, the development of hummingbird pollination must have followed one of two paths: (1) if once there were insect-pollinated species of Gesneria, they are now no longer extant and may have declined in response to more-effective methods of pollination; or (2) Gesneria as a genus evolved after the mech- anism for hummingbird pollination had been established in the family. Hummingbird pollination is well known in the Gesneriaceae (Stearn, 1969), so the possibility of hummingbird pollination should be expected in Gesneria. In fact, Harborne (1966) mentions a NUMBER 29 29 TABLE 4.-Characters of flowers visited by hummingbirds and bats* Flower character I ilumingbird$ Bats An thesis Color Corolla Odor Nectar guides Nectar availability Nectar quantity P o l l e n quantity diurnal vivid, usually red with contrasting Colors strong, tubular with no lip none none hidden in base of corolla tube or in spurs abundant moderate nocturnal or crepuscular drab, white, Cream to green or purple strong single flowers with large mouths or inflorescences of small flowers stale or sweet, or of ripe or decaying fruit none accessible abundant abundant Compiled from Faegri & van der Pijl (1966, 1971), Baker (1961), Meeuse (1961), and Grant L Grant (1968). report of hummingbirds visiting a Gesneria species. Of the 15 species of hummingbirds reported by Bond (1961) from the West Indies, 13 are selected here to be potential pollinators of Gesneria spe- cies. In Table 5 are listed the hummingbird species, the mean lengths of their bills, and one of the species of Gesneria which has a corolla tube length corresponding to the bill length. Hum- mingbird species are illustrated in Figures 21 and 25 with flowers of species they may visit and pollinate. The potential pollinators for some of the species will be discussed under the species. Sections Pentarhaphia and Physcophyl lon in par- ticular have the syndrome of characters found in flowers usually visited by hummingbirds. The color of the corolla tube ranges from pure red to light yellow with a red limb or greenish-white with a reddish base. The pigments of two Gesneria species are similar to the anthocyanidins in other bird-pollinated species in the Gesneriaceae (Har- borne, 1966). The anthocyanidins in the Ges- neriaceae are unique in the flowering plants and found elsewhere only in a few fern species. The corollas also differ from each other in size and shape (Figures 16d-k, 21a), but the flowers often have a horizontal orientation, which seems geneti- cally controlled, at least in Gesneria pedicellaris. This species grows on vertical cliffs in the Dominican Republic. I n such a habitat corollas are oriented perpendicularly to the substrate (Figure If); however, when grown in a pot in the greenhouse, the perpendicular orientation is main- tained, i.e., the corollas are oriented vertically. Stamens and stigmata are usually in a position to contact the hummingbird at the upper side of the bill base or on the front of the head. Since the stigma is receptive one or two days before or after the pollen is released, there is minimal chance of self-pollination. Since hummingbirds are strongly territorial in their habits (Stearn, 1969) and restrict their feeding to a limited area, populations of hummingbird- pollinated species of Gesneria may become inbred and form apparent endemic species. Humming- birds, however, are not flower-constant; thus if more than one potentially cross-breeding population of Gesnerieae occurs within the feeding area, nat- ural hybrids may result, e.g., Gesneria scabra var. sphaerocarpa X R h y t i d o p h y l l u m tomentosum, and Gesneria acaulis var. glabrata X R h y t i d o p h y l l u m t o m e n t o s u m (Figure lj), which have been discov- ered in Jamaica. The second type of vertebrate pollination in- volves bats. This facet of pollination biology has been neglected until recently, perhaps because of the nocturnal habits of bats, the flowering habit of the flowers visited by bats, and the restriction of this phenomenon to tropical regions. The first suggestion that bats might be pollination agents is attributed to Moseley (1879), although the first record of bats visiting flowers was made by De La Nux in a letter to Buffon in 1772 (Baker and Harris, 1957). In the past decade many additional observations of bat pollination have been made, as well as detailed studies of mechanisms of bat pollination (Vogel, 1969a, 196913). Nectar- and pollen-feeding bats in the New World belong to the suborder Microchiroptera of the order Chiroptera and represent two sub- families, Glossophaginae and Phyllonycterinae, of the family Phyllostomidae. These small mammals have heads with elongated narrow rostra and jaws that can fit into large campanulate flowers. Brush- like papillae near the tip of their extensile tongues are adapted for lapping nectar and pollen, allow- ing the bats to reach the nectar at the base of the corolla (Wille, 1954). According to Allen (1939), ?bats are guided by their sense of smell in finding the blossoms and are attracted by the nectar.? The odor, however, may not necessarily be detectable by humans. Kaisila 30 SMITHSONIAN CONTRIBUTIONS T O BOTANY TABLE 5.-Potential hummingbird pollinators (Trochilidae) and Gesneria species in the West Indies Island Cuba Hispaniola Jamaica Puerto Rico Lesser Antilles Hummingbird Species Mean bill length Gesneria species Mean corolla tube length (in cm) (in cm) Archilochus colubris 2.0 G. cubensis Chlarostilbon ricordii Calypte helenae Anthracothorax dominicus Chlorostilbon swainsonii Xellisuga minima Anthracothorax mango Mellisuga minima Trochilus polytmus Anthracathorax dominicus Anthracothorax viridis Chlorostilbon maugaeus Orthorhynchus cristatus Sericotes holosericeus Glaucis hirsuta 1 . 7 1.1 2.4 1.9 1.2 2.6 1.2 2.1 2.4 2.7 1.5 1 . 4 2.2 3.1 G. duchartreoides G. humilis G. pedicellaris ci. decapleura G. pulverulenta 6. acaulis G. scabra var. scabra G. scabra VBT. fawcettii G. cuneifolia G. cuneifolia G. citrina G. citrina G. pauciflora. G. ventricosa 1.8 1.4 1.3 2.5 2.2 1.3 2.5 1.2 1.5 2.5 2.5 l . R 1 . 8 2.2 2.6 (1966) reports a bat, Rousettus aegyptiacus, as the only visitor to the apparently odorless Bombax malabaricum. In flower-feeding bats the olfactory lobes of the brain are well developed; Moller (1932) has shown these organs to be far better developed in the nectar-feeding Glossophaga than in insectivorous bats. Some bats (e.g., Glossophaga, Brachyphylla) are able to hover and feed on the flower. Other bats (e.g., Leptonycteris nivalis in Arizona) cannot hover, but must make repeated feedings at a single flower. This they accomplish by flying up to a flower, stopping suddenly, lapping nectar and flying off to the side before falling from the flower (Cockrum and Hayward, 1962; Novick, 1969). Chiropterophilous flowers display a syndrome of characters distinctly different from those of flowers visited by insects and hummingbirds (see Table 4). Preadaptation of flowers and bats for coevolu- tion was discussed by Vogel (1969a, 1969b), who inferred that flowers adapted for bat pollination were present before the bats. Unless one accepts preadaptation, one can assume that plants adapted for bat pollination evolved from a bird-pollinated ancestor simultancously with the flower-feeding bats. The bats probably evolved from insect- feeding forms. Grant and Grant (1968:94) wrote that bat pollination may have emerged because hummingbird pollination in some tropical regions reached a saturation point ?conducive to the emerg- ence of a new derivative pollination system.? The idea of bat pollination in Gesneria was first proposed in 1969 by Vogel for G. viridiflora and G. calycosa. Although Vogel presented no definite proof, he based his hypothesis on flower structure and color. The species of Gesneria suspected of being bat pollinated display the characters listed in Table 4. The flowers are simi- lar to Kigelia, a member of the Bignoniaceae, which has been observed to be pollinated by bats in the Old World (Harris and Baker, 1958; Meeuse, 1961). The flowers are campanulate (Figures lh; 16b,c,m; 22a,b), borne on long erect or pendent peduncles, e.g., Rhytidophyllurn grande (Figure lc) or Gesneria pedunculosa (Figure li). No floral odor is apparent, but the resin produced by Gesneria sometimes has a fruity odor which may act as the attractant. Abundant nectar nearly fills the drab-colored to green corolla. The stamens and stigma appear oriented to touch the top of the head, rostrum, or nose leaf of the animal (Figure 22a,b). Much more pollen is produced by these species than by those pollinated by birds. One or two anthers of Gesncriu calycosa (probably bat pollinated) con- tain as much pollen as about 50 anthers of G. NUMBER 29 31 acaulis (probably hummingbird pollinated). The stigma is enlarged and may make greater contact with the bat?s head. The habit of a bat-pollinated plant is usually fruticose or arborescent, bringing the flowers high above the ground where bats commonly fly. Re- ports of flower-feeding bats, Monophyllus redmani, being observed to ?settle on walls low down? (Osburn, 1865) or caught in ground-level nets (Tamsitt, 1967) suggest that some bats are able to visit flowers at or near ground level. Gesneria pumila and other species of short stature commonly grow on rock ledges, making the plants as accessible to flying mammals as those with a taller habit. Eighteen species and subspecies of flower- FIGURE 22.-Bats with flowers they may visit and pollinate: a, specimen of Brachyphylla cavernarum posed with Gesneria calycosa to show relative size of bat?s head and flower (note contact of anthers with top of bat?s head): b, specimen of GlossoPhaga soricina ant i l larum posed with Xhytidophyllurn tonentosun (arrow points to contact of nose leaf with anthers). feeding bats are known from the West Indies (Hall and Kelson, 1959; Schwartz and Jones, 1967; McNab, 1971); from these, fifteen may be potential pollinators for Gesneria species. I n Table 6 there are listed potential bat pollinators from each of the islands, critical measurements of bat heads, and species of Gesneria which have corollas corresponding to the size and shape of the bat head. Two species of bats are illustrated in Figure 22a,b. All of the major islands of the Greater Antilles have species of Gesnerin that could be bat pol- linated. Of the four largest islands, Jamaica has the largest number of these forms, as well as the largest number of extant bat species. This fact may reflect the opinion of Koopman and Williams (1951) that Jamaica. was ?a way station on the road to invasion of the West Indies? and that in the Greater Antilles bat pollination may have developed first in Jamaica and spread to the other islands by land bridges in the geologic past. In a later article, Koopman (1958) suggests that the poorer total representation of bats in the Greater Antilles, as compared to the American continents and their nearby islands, is the result of a combina- tion of water gaps and environmental restriction. Although most of the Antilles have been above sea level since the Late Miocene-Early Pliocene (Schuchert, 1935), Walker (1968) states that the family Phyllostomidae a2peared no earlier than the Pleistocene. According to Allen (1939), fossil deposits containing remains of bats in the West Indies are no older than the late Pleistocene, although fossil bats can be found in European deposits dating back to the Eocene. Some members of the flower-feeding bat genera are now known only as subfossils. Allen suggests that the disap- pearance of some of the taxa within the past cen- tury or more may be due to human habitation and forest destruction. Pollen dissemination in Gesnerin is more likely dependent on the advanced mechanisms of hum- mingbird and bat pollination to achieve cross- breeding within populations than upon pollina- tion by insects. Evidence to prove this hypothesis can be found in the characters of these tropical organisms. A1 though evidence has been given to show the likelihood of hummingbird and bat pollination, only direct observation can prove the existence of such mechanisms. 32 SMITHSONIAN CONTRIBUTIONS T O BOTAKY Island Cuba Hispaniola Jamaica Puerto R i m TABLE 6.-Potential bat pollinators (Phyllostomidae) and Gesneria species in the West Indies Bat species and subspecies Mean head Mean rostrum Gesneria species Corolla tube width at eyes length: eye (means, in cm) Width Length (LP. cm) to noseleaf (lr. m) (at mouth) Brachyphylla nana Erophylla sezekorni sezekarni Monophyllus redmani clinedaphus Phyllonycteris poeyi Brachyphylla pumila Erophylla bombifrans santacristobalensis Monophyllus redmani clinedaphus Erophylla sezekami syaps Brachyphylla pumila Glassophaga soricina antillarum Monophyllus redmani redmani Phyllonycteris aphylle Brachyphylla cavernarum Erophylla bombifrans bombifrons Monophyllus redmani partoricensis 1.0 0.8 0.8 1.0 1.2 0.8 0.8 0.8 1.2 0 . 9 0.8 1.3 1.3 0.8 0 .6 1.1 0 . 9 0 . 9 0.8 1.0 0.8 0 . 9 0.9 1.0 0.8 0 . 9 1.1 1.2 0.8 0.8 G. shaferi G. viridiflora subsp. viridiflora G. gloxinioides G. viridiflora subsp. viridiflora G. fruticosa 6. viridiflora subsp. quisqueyana G. viridiflora mbsp. quisqueyana G. pumila subsp. mimuloides G. clandestina G. exserta G. pumila subsp. neglecta G. calycosa G. viridiflora subsp. sintenisii 6. pedunculosn G. pedunculosa 1.0 1.0 0.8 1.0 1.8 1.0 1.0 0 . 7 2.0 1.2 0 . 7 1.4 1.2 1.0 1.0 1.1 1.2 1.3 1.2 1.8 1.2 1 .2 1.2 2.1 1.5 1.2 1.8 1.2 1.5 1.5 METHODS OF SEED DISPERSAL IN Gesneria Coevolution for the dispersal of fruits and seeds has occurred among many plants and animals (Stebbins, 1971), although perhaps not to any great degree in Gesneria. T h e fruit is a dry capsule containing very small, numerous, reddish-brown to black, twisted seeds. Three methods for dis- persing the seeds of Gesneria may be important. 1. Transport by animals. Large animals would be only incidental carriers of seeds, but inverte- brates may be important. Because ants find the seeds of other members of the Gesneriaceae attrac- tive and may aid in dissemination (Stearn, 1969), myrmecochory or dispersal by ants is suspected. Stebbins (1971) enumerated the structural modi- fications found in ant-dispersed species: (a) the inflorescences are near the ground. This character is common among the acaulescent spccies, such as Gesneria acaulis, G. cuneifolia, G. pumila, and others; (b) the peduncle is recurved at maturation. This arching of the peduncle places the capsule near the ground and is noticeable particularly in Gesneria pumila subsp. neglecta; (c) the capsules dehisce irregularly over a long period of time; and (d) the seeds may bear an elaiosome or fat- bearing appendage. This last structural modifica- tion is not well developed in Gesneria, but the funicle of thc seed is occasionally enlarged or elongated (Figure 19d,f,h) and may be attractive to ants. Depcnding upon the adaptations it seems likely that for acaulescent species of Gesneria growing in tropical forests, myrmecochory may be a major means of dispersal. Stebbins (1971) con- cludes that for herbs in forests, ant dispersal is one of the most effective means of dissemination. 2. Dispcrsal by water. Plants of Gesneria reti- culata, G. humilis (Figure la), and G. pauciflora usually grow in or near running water and often show signs of inundation. If a plant with open capsules were submcrged or shaken to allow the seeds to fall on the surface of the water, the seeds might bc carried downstream to a new locality. Not only seeds, but also whole plants caught in a floating mat of vegetation might be distributed NUMBER 29 33 this way. Capsules of the species mentioned above dehisce so that the open fruit is oriented vertically to become a splash cup, a structure heretofore found only in a few flowering plants, e.g., Mitel la (Saxifragaceae) and Sagina (Caryophyl1aceae)- but analogous to the gemmae cups in Marchant ia and to structures that function in the dissemination of spores in the Nidulariales or bird?s nest fungi. During heavy rains or near waterfalls, drops of water may fall into the capsule with such force as to eject the seeds, a phenomenon called ?rain- ballists? by van der Pijl (1969). In the splash of a waterfall in the Dominican Republic, I observed plants of Gesneria reticulata with this type of dis- persal mechanism operating. One plant had within its open capsules seedlings that had germinated in the humid atmosphere and were being washed out of the capsule by the spray of the waterfall and carried downstream. 3. Dispersal by wind, This method is especially important for those species that grow on dry cliffs or at forest margins. The dry capsules dehisce from the apex and often invert during maturation to spill out the seeds like a salt shaker, the ?censer- mechanism? (van der Pijl, 1969). The capsules have enlarged ribs or costae that catch currents of air to shake the capsules, sifting out the seeds. The wide persistent sepals also may catch air currents. A wind strong enough to move the capsule would likely carry the seeds as ?wind-ballists? for some distance from the parent plant. The wind-dispersal method may explain in part long-distance distribution of species in the tribe Gesnerieae. Such dispersal over a wide geographic range is not common; only a few species range over more than one island: i.e., Gesneria reticulata (Puerto Rico, Hispaniola, and Cuba); Gesneria humi l i s (Hispaniola and Cuba); R h y t i d o p h y l l u m t o m e n t o s u m (Cuba and Jamaica): Rhyt ic lophyl lum aur icu la tum (Puerto Rico and Hispaniola). There are, in addition, a number of groups of closely related species distributed among the islands: e.g., Gesneria christii in Haiti, G. acaulis in Jamaica, and G. purpurascens in Cuba; G. p u m i l a in Jamaica and G. shaferi subsp. depressa in Cuba; G. pedicellaris in the Dominican Republic and G. libanensis in Cuba, These distributions may be explained in either of two different ways: (1) The plants we see today may be relicts of a more extensive range in the geologic past. This seems unlikely, for the species appear to be relatively young in geologic time and undergoing relatively rapid evolution. (2) The range existing at the present time may be due to transport by air currents in recent times by tropi- cal storms or hurricanes. Stebbins (1971) states, ?the incidence of strong winds is erratic and unpredictable, so that they may or may not occur when seeds are ripe and ready to be transported. Furthermore, the direction of winds is variable and they tend to scatter seeds over a large territory, including both favorable and unfavorable habi- tats.? Hurricanes occur in the West Indies with annual regularity and their tracks are usually in a northwesterly direction. Stebbins also notes that if the fruit and seeds are very small, long distances can be covered from the parent plant. And such is the case in the Gesnerieae, The small seeds of Gesneria reticulata, for example, could possibly travel the air currents of a tropical storm from Puerto Rico to Hispaniola and then to Cuba. Long-distance transport of seeds by wind may occur more frequently than we realize, but the results are not evident due to the limited success of the immigrant plants whose environmental re- quirements are not being met in the new habitats. According to Ashton (1969), ?In the tropics . , ., ground herbs appear little adapted to environ- mental, at least edaphic, variation, though isola- tion of small morphologically distinct populations is common in many groups, as the Gesneriaceae.? Of the three dispersal mechanisms suggested- transport by animals, water, and wind-the first two are likely means for short-distance dispersal, but the third method is the only one feasible for long-distance distribution. Wind dispersal, if coupled with a possible capacity for self-pollination and environmental adaptation, may account for the distribution of some of the species of Gesneria. Hybridization and Cytology Cytology reveals chromosomal characters within the cell, while hybrid studies may reveal genetic relationships not apparent in the parent species or may identify barriers that prevent hybrid produc- tion. Cytological studies in the Gesnerieae will be reviewed in relation to hybridization experiments, and mechanisms will be suggested that may account for evolution in the tribe. 34 SMITHSONIAN CONTRIBUTIONS TO BOTANY MATERIALS AND METHODS Living plants for study were obtained from many sources and added to the collection of Gesneriaceae already in cultivation in the green- houses of the L. H. Bailey Hortorium: (1) seeds and cuttings were obtained during a plant col- lecting trip to the West Indies in 1970; (2) cuttings were received from the living collections at the Royal Botanic Garden, Edinburgh, Scotland; the Royal Botanic Gardens, Kew, England; Fairchild Tropical Garden, Miami, Florida; and from the U.S. National Arboretum, Washington, D.C.; (3) seeds and plants were obtained from the seed fund and members of the American Gloxinia and Gesneriad Society. Voucher specimens of materials are filed in the herbarium of the L. H. Bailey Hortorium, Cornell University. Environmental factors play a large role in the success of species and hybrids in the greenhouse or in nature. T o attempt to find the ideal conditions or to simulate the natural conditions, a variety of temperature, substrate, and humidity regimes was tried. Plants were grown in the greenhouse at the L. H. Bailey Hortorium in each of two tempera- ture ranges (55-70?, 65-75O). A few plants were also grown in a greenhouse at Guterman Bio- climatic Laboratories of Cornell University where they received a higher light intensity and lower humidity. The standard substrate was ?Cornell Epiphytic Mix? formulated by R. Mott (1972). Because many of the species in the wild grow on calcareous soil, calcium nitrate and chelated iron were added occasionally to the substrate (Skog, 1972b). Crosses were made in the greenhouse using fresh pollen or stored pollen prepared in the method recommended by Clayberg (1964). Pollen viability was tested by staining a small quantity of fresh pollen on a slide with either cotton blue in lacto- phenol or acetocarmine. (The latter stain used by R. E. Lee proved to be more reliable.) One thou- sand grains were counted to determine percent stainability. Seeds developed in two to five months. Because of the suspected poor viability of hybrid seed, germination was attempted soon after dehiscence of the capsule. RESULTS OF HYBRIDIZATION EXPERIMENTS Artificial hybrids can be obtained with relative ease in the Gesnerieae under cultivation. Appar- ently active isolation mechanisms prevent wide- spread hybridization in the wild. Putative natural hybrids do occur, however, in rare instances. Two natural hybrids have been collected in Jamaica and brought into cultivation (Gesneria scabra var. sphaerocarpa X Rhytidophyllum tomentosum and Gesneria acaulis var. glabrata X Rhytidophyllum tomenlosum). The latter hybrid has been collected in a few different localities in Jamaica and was considered a distinct species in Gesneria by Adams (1972). In the greenhouse I attempted about 240 crosses, including intergeneric crosses between Gesneria and Rhytidophyllum, interspecific crosses within each genus, backcrosses of the hybrids to the par- ents, reciprocal crosses by alternating pollen and seed parents, and crosses between hybrids. Only 12 crosses resulted in viable seed, producing plants that survive today. These plants were added to the stock of hybrids produced by Wiehler, Katzenberger, and others already in cultivation in the greenhouses at Cornell University. Twenty- nine hybrid forms are now known. The most recent crosses have not yet flowered; statements made about morphology are derived from the few that have flowered. Crosses attempted and results ob- tained are listed in Tables 7 and 8. In general, interspecific hybrids at the F1 stage are intermediate in morphology between parents; however, intergeneric crosses reveal the following: 1. Crosses between plants with campanulate corollas and those with nearly cylindric corollas result in hybrids with nearly cylindric corollas (Figure li), implying that genes for the nearly cylindric corolla dominate. Crosses between two species with campanulate corollas, e.g., Gesneria fruticosa X Rhytidophyllurn tomentosum yield forms with a campanulate corolla but intermediate in color and vestiture. 2. Offspring are, in general, more vigorous in growth, taller, and more erect than either parent. During the extended flowering season more flowers are produced. Hybrids, however, appear to be more sensitive to some environmental stresses than the parents, suggesting inherent hybrid weakness. 3. Leaf shapes of intergeneric hybrids tend to NUMBER 29 TABLE 'i.-Hybrids and parentage 35 TABLE 8.-UnsuccessfuI crosses Hybrid 6-number 1240 1230 1238 1237 1236 1235 1241 1249 1239 1245 1231 1406 1405 1233 1418 1421 1420 1036* 1402 1404 1419 1403 1398 1232 1234 1401 1399 1371* 138411 - Seed Parent Pollen Parent G-number name 6-number name 876 876 877 888 888 888 857 897 898 898 898 767 767 767 767 767 1034 1256 1256 1257 1257 1257 1010 1010 841 1381 Gesneria acaulis var. glabrata 6. acaulis VBT. glabrata 6. acaulis var. acaulis 6. citrina G. citrina G. citrina 6. cuneifolia 6. reticuiata 6. pedicellaris G. pedicellaris G. pedicellaris G. pedunculosa G. pedunculosa 6 . pedunculosa 6. pedunculasa G. pedunculosa G. pulverulenta G. scabra var. sphaerocarpa Rhytidophyllum auriculatum R. auriculatum R. berteroanum R. berteroanum R. berteroanum R. leucomallon R. leucomallon R. tomentosum R. tomentosum R. tomentosum R. tomentosum 940 908 940 876 767 940 940 908 767 942 883 1037 940 765 1257 883 1226 1257 1260 767 905 841 767 883 767 1256 Gesneria ventricosa subsp. ventricasa Rhytidophyllum auriculatum G. ventricosa subsp. ventricosa G. acaulis var. glabrata G. pedunculosa 6. ventricosa subsp. G . ventricasa subsp. R. auriculatum 6. pedunculosa G. viridiflora subsp. quisqueyana R. grande 6. calycosa G. ventricosa subsp. ventricosa R. auriculatum R. berteroanum R . grande G. pedicellaris R. tOmentOSum R. berteroanum R. auriculatum G. pedunculosa R. auriculatum R. tomentosum G. pedunculose R. grande G. pedunculosa R. auriculatum G. acaulis G. fruticosa ventricosa ventricosa * = putative natural hybrid. # = cross made by R. Katzenberger. resemble more those of the Rhytidophyllum parent, i.e., narrowly Ianceolate, very elongated, areolate with prominent veins. 4. The numerous inflorescences far exceed in length the subtending leaves, a common trait in Rhytidophyllum. 5. Internodes are elongated even more than the normally elongate internodes in Rhytidophyl- lum, but are less hirsute and with the erumpent lenticels found in Gesneria. 6. Pollen stainability, a test for viability and fertility, is very low, from less than 1 percent (Gesneria acaulis var. glabrata X Rhytidophyllum tomentosum) to 35 percent (Gesneria scabra var. sphaerocarpa X Rhytidophyllum tomentosum). Pol- len stainability in interspecific hybrids ranges from 0-66 percent. At the present time neither backcrosses to parents nor crosses between hybrids have produced seeds, suggesting that low-pollen viability or chromosomal anomalies may prevent fertilization. In vegetative morphology, reciprocal crosses appear similar; none have yet flowered. Crosses with Gesneria Potential seed Potential pollen parents, G-number* parent. G-nurnber* 763 767 769 784 841 857 869 876 877 881 883 888 897 898 903 905 907 908 940 942 1008 1034 1036 1037 1066 1226 1231 1235 1236 1238 1239 1240 1244 1245 1249 1256 1257 1260 1268 1360 1361 1365 1381 767, 1037 784, 841, 1034 767, 841, 1034, 1037 ( 2 ) , 1256, 1257, 1381 767, 1037, 1257 960, 1036, 1037, 1256, 1257 767. 881 767, 1037 767, 841, 883, 1037, 1256 (2), 1361, 1381 RL1 . _ 767, 841, 1256 841, 876, 1034, 1037, 1256, 1257, 1381 1037, 1381 1381 1034 (2), 1037, 1245, 1256, 1381 940, 1037 767, 883, 940, 1037 (2) 1037 1257, 1260 767. 841. 883 903' 1 n 3 i ... 767 (2), 769, 883, 897 (2), 940, 1037 (2), 1256, 1257 (2), 1260, 1365, 1380 841, 1037, 1257, 1260 767. 841 ( 2 ) . 876, 883, 903, 905, 1257, 1268 841, 903, 905 767 (2), 841, 881 (21, 940, 1037 (2) RL1 i6i, 1256 767 841 767, 1256 876, 940 767 7 6 7 . 898 . , ~.~ 897, 1037, 1260 767 (2), 769, 841, 883, 1034, 1037, 1260 767, 876, 883 (2), 903, 940, 1034, 1037, 1256, 1365, 1381 767, 940, 1037, 1256 841 (2), 1037, 1257 RL1 .. 767, 1037 765, 767 (Z), 769 (2), 784, 841, 876 (2), 883, 897, 940, 1034 (41, 1036, 1037 (3), 1226, 1256, 1257 767, 841, 876, 940, 1034, 1037, 1260 * Refer to Table 9 far s p e c m IWIC 01' L-nmmcr. ( ) = number of attempts mare than one. humilis or Pheidonocarpa corymbosa as either the pollen or seed parent have failed. DISCUSSION OF CYTOLOGY AND HYBRIDIZATION Until 1969 a haploid number of 14 chromosomes was believed to be the base number in the Ges- nerieae; however, Nevling (1969) reported a count of n = 7 in a plant of Gesneria viridiflora subsp. sintenisii. This report leads one to suspect that seven chromosomes in each gamete is the basic number and that those plants with a haploid number of 14 are tetraploids. Polyploids are considered rare in the American members of the Gesneriaceae (Wiehler, 1972); only 18 tetraploids have been determined from 193 species in 37 genera. In flowering plants as a group, however, polyploidy is believed to occur in 30-35 percent of the species (Stebbins, 1950) or up to 43 percent (Grant, 1963). The first chromosome count of a plant in the 36 SMITHSONIAN CONTRIBUTIONS T O BOTANY Gesnerieae was made in 1956 by Eberle, who demonstrated that in Gesneria tomentosa L. (=Rhytidophyllum tomentosum) n = 14. Since then chromosome counts of many species have been determined by R. E. Lee (1964, 1966a, 196613, 1968). The only other number differing from n = 14 was n = 28 in a collection of Gesneria cuneifolia, which Lee (1964) considered a tetraploid. Grant (1963) suggested that a haploid number of 7, 8, or 9 chromosomes is basic in angiosperms. He concluded that plants with a gametic number of 14 or more may be considered polyploids. In the Gesneriaceae, counts of n = 8 or n = 9 have been reported many times, as well as a few counts of n = 16 or n = 18 and others, but n = 7 is known at the present time only from Gesneria uiridiflora subsp. sintenisii. According to Grant (1971) the promotion of polyploidy in a group of plants results from a combination of factors: (1) the plants are ?long- lived organisms usually possessing means of vege- tative propagation?; ( 2 ) original speciation in the genera was accompanied by a repatterning of the genes in the chromosome; and (3) natural inter- specific hybridization is a common occurrence. Plants in the Gesnerieae fit the first factor, for they are woody-stemmed plants apparently capable of surviving for many years and propagating vegeta- tively. The second factor is unknown in Ges- neriaceae and cannot be determined without detailed knowledge of the gene patterns in chromosomes. Interspecific hybridization, however, is likely. Since only putative tetraploid artificial interspecific hybrids are known, one cannot judge the relative occurrence of natural diploid inter- specific hybrids. Perhaps, these hybrids resembled one of the parents to such a degree that they were indistinguishable in the diploid condition. Pos- sibly only after doubling of the chromosomes did the differences become evident. Interspecific dip- loid hybrids may have occurred with considerable frequency in the past; one can assume that if hybridization can occur at all, it should have occurred frequently relative to geologic time. If two intergeneric hybrids (which may be produced with more difficulty than hybrids between species in the same genus) can occur in our time and be seen readily, interspecific hybrids should have been readily obtainable in the past. According to Lewis (1966), in some members of the Scrophulariaceae the diploid species tend to be self-incompatible and the natural polyploid species self-compatible. If this is also true in the Ges- neriaceae and in the tribe Gesnerieae, the poly- ploid species may have had reproductive advantage over the diploids and because of heterosis may have been ecologically superior and able to spread to niches not open to diploid species. Stebbins (1950) writes that polyploids are well adapted for colonizing newly exposed habitats. Observation of plants in the field suggests that most Rhytido- phyllurn species and perhaps Gesneria exserta and G. pedunculo.ta do adapt to new areas readily. Systematic Treatment The tribe Gesnerieae consists of approximately 67 species in three genera, of which Gesneria and Pheidonocarpa will be treated in detail here. For completeness, however, a key is given to distin- guish the genera within the tribe. The 46 herbaria whose collections were made available to me are abbreviated as given by Holmgren and Keuken (1974), except for one not included in the sixth edition of Index Her- bariorum, RD J-private herbarium of Dr. Jose Jimenez, Santiago de 10s Caballeros, Dominican Republic. Tribe GESNERIEAE Tribe GESNERIEAE, G. Don, Gen. Syst. 4:643-644, 1838.-DC., Prodr. 7:525, 1839.-Fritsch in Engl. & Prantl, Nat. Pflan- zenfam. 4 (3b):183, 1894. GUNERIEAE subtribe CONRADIEAE G. Don, Gen. Syst. 4:644, 1838. [Type-genus: Conradia Martius.] EUGESNEREAE Endlicher, Gen. P1. 721, 1839. GESNEREAE subtribe RHYTIDOPIIYLLEAE Hanstein, Linnaea 26: 199, 204-205, 1854 [pro parte]. [Type-genus: Rhytidofihyl- lum Martius.] RHYTIDOPHYLLEAE Orsted, Cent. Gesn. 10, 1858 [?Rytidophyl- leae?] Hanst., Mart. F1. Bras, 8 (1):394, 1864. [Type-genus: Rhytidophyllutn Martius.] GESNEREAE subtribe PENTARHAPHIEAE Bentham in Bentham & Hooker, Gen. P1. 2:993, 1876. [Type-genus: Pentarhafihia Lindley.] Fibrous rooted, acaulescent perennial herbs, sub- shrubs to trees: stems woody, usually resinous. Leaves alternate and spiral or opposite and decussate, sessile or petiolate, ?pseudo-stipules? NUMBER 29 G-number Taxon Area of Origin 37 G-number T@.XO" Area of Origin 763 765 767 769 784 841 857 869 874 875 876 877 881 883 888 897 898 901 902 903 905 907 908 909 924 940 942 1008 1010 1034 1035 1036 1037 1066 1067 1226 1230 1231 1232 1233 1234 1235 1236 1237 1238 1239 1240 1241 1244 1245 Gesnerie cuneifoiie Rhytidophyllum auriculatum G. peduncuiosa G. pauciflora G . reticulata R. tomentosum G. cuneifolis G. cuneifolie G. pumila subsp. pumila G . pumila subsp. neglecta G. acaulis var. glabrata G. acaulis var. acaulis G . scabra var. sphaerocarpa R. grande G. citrina G. reticulata G. pedicellaris G . exserta G. viridiflora s v b e p . sintenisii G . viridiflora subsp. quisqueyana R. auriculatum R. auriculatum R. auriculatum R. auriculatum R. auriculatum G. ventricasa rubsp. ventricosa G. viridiflora subs?. quisqueyana G. christii R. leucomallon G. pulverulenta G . fruticosa G. scabra VBT. sphaerocarpa X R. tamentosum G. ca1ycosa R. auriculatum R. auriculatum G . pedicellaris G. acaulis var. glabrata (876) G. pedicellaris (898) x R. grande (883) R. leucomallon (1010) G. pedunculosa (767) R. levcornallan (1010) X R. grande (883) G. citrina (888) X G. ventricosa aubsp. X R. auriculatum (908) X G . peduncuiosa (767) X R, auriculatum (765) ventricosa (940: G. citrina (88 '8) X G. pedunculosa (767) G. citrina (888) X G . acaulis (876) G . acaulis var. acaulis (877) G . pedicellaris (898) X G . pedunculosa (767) G. acaulis var. clabrata (876) X G . ventricosa subsp. ventricosa (940) Puerto Rico Puerto R i m Puerto Rico Puerto Rica Puerto Rico Jamaica Puerto Rica Puerto R i m Jamaica Jamaica Jamaica Jamaica Jamaica Jamaica Puerto R i m Dominican Republic Dominican Republic Jamaica Puerto Rico Dominican Republic Dominican Republic Dominican Republic Dominican Republic Dominican Republic Haiti St. Kitts Dominican Republic Haiti Haiti Dominican Republic Dominican Republic Jamaica Jamaica Dominican Republic 7 Dominican Republic * * , * X G . ventricasa subsp. ventricosa (940) * X G. ventricosa subsp. ventricosa (940) G . cuneifolia (857) G . cuneifolia G. pedicellaris (898) X G . viridiflora subsp. quisqueyana (942) * 1248 1249 1256 1257 1260 1265 1268 1355 1360 1361 1362 1363 1364 1365 1366 1367 1368 1369 1370 1371 1372 1373 1374 1375 1379 1380 1381 1384 1385 1395 1398 1399 1400 1401 1402 1403 1404 1405 1406 1418 1419 1420 1421 1453 1454 1476 G. vencricosa subsp. ventricasa G. reticulata (897) X R. auriculatum (908) R. auriculatum R. berteroanum R . auriculatum R. auriculatum R. auriculatum R. leucomallon G. cuneifolia G. pauciflora G . viridiflora subsp. sinteniaii G. ventricosa subsp. ventricosa G. pedicellaris G. humilis G. haitiensis Pheidonocarpa corymbosa G . clandestina G. exserta G. harrisii R. tomentosum X G . acaulis R. auriculatum R. berteroanum G. reticulata R. auriculatum G. fruticosa G . pumila subsp. pumila R. tomentosum R. tomentosum X G. fruticosa R. tomentosum G. pedicellaris R. berteroanum (1257) R. tomentosum (1381) G. jamaicensis R . tomentosum (841) X G. pedunculosa (767) R . auriculatum (1256) X R. berteroanum (1257) R. berteroanum (1257) X R. auriculatum (905) R . auriculatum (1256) X R . auriculatum (1260) G. pedunculosa (767) X G. ventricosa subsp. ventricosa (940) G. pedunculosa (767) X G. calycosa (1037) G . pedunculosa (767) X R. berteroanum (1257) R. berteraanum (1257) X G. pedunculosa (767) G. pulverulenta (1034) X G. pedicellaris (1226) G. pedunculosa (767) X R. grande ( 8 8 3 ) G. viridiflara subsp. quisqueyana G. pumila subsp. pumila R. auriculatum X R . tomentosum (841) X R. auricultltum (1256) t Dominican Republic Dominican Republic Dominican Republic Dominican Republic Dominican Republic Puerto Rico Puereo Rico Puerto Rico Dominica Dominican Republic Haiti Haiti Jamaica Jamaica Jamaica Jamaica Jamaica Dominican Republic Dominican Republic Dominican Republic Dominican Republic Haiti Jamaica cross by R . Katienberger Jamaica 7 * Jamaica * * * * * * * * Dominican Republic * = hybrid of greenhouse origin. ? = plant obtained from horticultural source, loceliry unknown. Gesneria acaulis var. acaulis 877 R. leucomelion 1010, 1355 G . acaulis var. glabrata 876 R. tomentosum 841, 1381, 1385 G . calycosa 1037 G. acaulis var. glabrata (876) X G. ventricosa subsp. ventrlcosa (940) 1240 G. christii 1008 G. acaulis "81.. glabrate (876) X R. auriculatum (908) 1230 G . citrina 888 G . scaulis var . acaulis (877) X G. ventricosa subsp. ventricasa (940) 1238 G. clandestina 1368 G . citrina (888) X G. acaulis var. glabrata (876) 1237 G. cuneifolia 763, 857, 869, 1244, 1360 G. citrina (888) X G . pedunculosa (767) 1236 G . exserte 901, 1369 G . citrina (888) X G. ventricosa subsp. ventricasa (940) 1235 G. fruticosa 1035, 1379 G. cuneifolia (857) X G. ventricosa subsp. ventricosa (940) 1241 G . haitiensis 1366 G. pedicellark (898) X G . peduncvlosa (767) 1239 G. harrisii 1370 G . pedicellaris (898) X G. viridiflora subsp. quisqueyana (942) 1245 G . humilis 1365 G. pedicellaris (898) X R. grande (883) 1231 G. jamaicensis 1400 G . pedunculosa (767) X G . ventricosa subsp. ventricosa (940) 1405 G . pauciflora 769, 1361 G . pedunculosa (767) x R. auriculatum (765) 1233 G . pedicellaris 898, 1226, 1364, 1395 G. pedunculosa (767) X R. berteroanum (1257) 1418 G. pedunculosa 767 G. pedunculosa (767) x R. grande (883) 1421 G. pulverulenta 1034 G. reticulate (897) X R. auriculatum (908) 1249 G . pumila subsp. neglecta 875 G . pulverulenta (1034) X G. pedicellaris (1226) 1420 G. pumila subsp. pumila 874, 1380, 1454 G. scabra Yar. sphaerocarpa X R. tomentosum 1036 G. reticulata 784, 897, 1374 R. auriculatum (1256) X R. auriculatum (1260) 1404 G. scabra var. sphaerocarpa 881 R. auriculatum (1256) X R. berteroanum (1257) 1402 G. ventricosa subsp. ventricosa 940, 1248, 1363 R. berteroanum (1257) X G. pedunculosa (767) 1419 G. viridiflora subsp. quisqueyana 903, 942, 1453 R. berteroanum (1257) X R. auricuiatum (905) 1403 G. viridiflara subsp. sintenisii 902, 1362 R. leucomalion (1010) X G. pedunculosa (767) 1232 Pheidonacarpa corymbosa 1367 R. leucamallan (1010) X R. grande (883) 1234 Rhytidaphyllum auriculatum 765, 905, 907, 908, 909, 924, 1066, 1067, 1256 1260, 1265, 1268, 1372, 1375, 1476 R. tomentosum (1381) X R. auriculatum (1256) 1399 R. berteroanum 1257, 1373 R. grande 883 R. tOmento8um (841) X G. pedunculosa (767) 1401 R. tomentosum X G. acaulis 1371 R. tomentosum X G . fruticoss 1384 38 S M I T H S O N I A N C O N T R I B U T I O N S T O BOTANY occasionally present; petioles with a trace nearly cylindrical or U-shaped in cross section; blades dorsiventral, usually with a multiple epidermis. Inflorescences modified compound dichasia or reduced to single flowers: peduncles elongated be- yond the leaves or shorter; floral tube of fused bases of sepals, petals and stamens adnate to the inferior or subinferior ovary, turbinate to sub- spherical; calyx with 5 lobes free or briefly united or rarely connate for about half their length above the ovary; corolla tube cylindrical to campanulate, red, orange, yellow, brown, green, or white; fertile stamens usually 4, rarely 5, staminode usually 1, all adnate to the base of the corolla tube; ovary of 2 carpels, uniloculate, ovules numerous, each with a single integument, disc annular, often 5-lobed. Capsule linear, obconic, turbinate or sub- globose, slightly raised apically, or rostrate and rostrum exceeding the length of the floral tube, capsule opening by 2 or 4 valves; seeds numerous, fusiform, rhombic or linear. TYPE-GENUS.-Gesneria Linnaeus. The concept of the tribe Gesnerieae has varied since its circumscription by George Don in 1838. Only after the circumscription by Fritsch in 1894, based on the characters of leaf arrangement, ovary position, and disc shape, was the tribe delimited to include Gesneria and Rhytidophyllum alone. In 1971, Wiehler transferred two species formerly in Heppiella to Gesneria, thereby including plants with opposite decussate leaves within the limits of the tribe. Since the two species appear to be more closely related to Gesneria and Rhytidophyllum than to any other genus or tribe, the limits of the Gesnerieae have been redefined. The tribe Gesnerieae is here composed of three genera whose limits are determined by characters of habit, flowers, fruit, and foliage. Parallel evolu- tion and adaptations to environment in all three genera may give the appearance of closer relation- ship than is real in nature. The corolla, in par- ticular, has apparently become adapted to pollinating agents, presumably bats and hum- mingbirds. Parallel adaptation to the same pollinating organisms has resulted in corollas of similar morphology within the three genera. Pheidonocarpa displays many morphological and anatomical features similar to Gesneria and Rhytidophyllum and has a gametic chromosome CHROMOSOME NUMBER.-n = 7, 14, or 28. number ( n = 14) common to many species of the other two genera. The genus, however, can be distinguished from Gesneria and Rhytidophyllum by its consistently opposite and decussate leaves and by its rostrate fruit. Pheidonocarpa resembles Rhytidophyllurn in its compound dichasial in- florescence, but probably did not evolve from that genus but frqm an ancestral stock of opposite- and decussate-leaved plants which had developed, like species of Gesneria and Rhytidophyllum, a chromosome number of n = 14. Gesneria and Rhytidophyllum appear closely related in floral characters, sometimes occupying similar niches and possibly utilizing the same pol- linators. The genera cannot be distinguished as separate units by any single factor, but by a com- plex of correlated characters, primarily those related to reproduction, but reinforced by vegeta- tive factors. Gesneria and Rhytidophyllum have traditionally been distinguished by the relative length of adnaiion of the filaments to the corolla tube; this trait does not hold in every case, but when correlated with foliage and fruit characters may determine the generic standing of a plant in these genera. The natural habitats of Rhytidophyllum, Ges- neria, and Pheidonocarpa may give clues to dis- tinguish the genera. In the field, Rhytidophyllum species usually occupy marginal land at roadsides (Figure Ib), waste places, cliffs, and forest margins where the plants are exposed to full sun and wind. Rhytidophyllum tornentosum in Jamaica, R. auri- culaturn in the Dominican Republic, and R . leucomallon in Haiti (Figure I b ) could be called weeds or pioneer plants, for they occupy newly exposed soil often to the exclusion of other plants. Species of Gesneria are usually restricted to more protected locations along streams or waterfalls, on ledges or clefts in cliffs or banks, or grow in damp forests as understory shrubs. Species which grow in clearings and/or roadsides and are exposed to full sun are located in damp soil or produce many branches from the base whose shade may maintain a lower temperature at the soil level. Gesneria citrina, which grows in Puerto Rico on barren limestone cliffs, has long roots that penetrate into the porous substrate and has dead leaves that persist and may protect the stem from the sun. The origin and evolution of the Gesnerieae is unknown. One may suggest an origin in northern NUMBER 29 39 South America from a woody-stemmed, fibrous- rooted, opposite-leaved, hummingbird-pollinated ancestral line, perhaps similar to Pheidonocarpa. One probable early derivative in South America, Gesneria onacaensis Rusby (= Rhytidophyllum), displays many of the primitive characters men- tioned above. From such a form two evolutionary lines may have developed in the directions of Gesneria and Rlzytidophyllum. Pheidonocarpa may have maintained characters similar to the ancestors of the tribe. T h e spread of the ancestral plants from northern South America may have occurred via the routes and mechanisms discussed under Pollination and Seed Dispersal. The close contact between the genera in their similar niches allows one to suspect occasional interchange of genes, through chance cross- pollination and fertilization and possibly rare back- crosses between parents and hybrid offspring. The possible interchange of genes is apparent in section Duchartrea of Gesneria. Members of this section resemble to a large degree a few species of Rhytido- phyllum (R. grande (Swartz) Mart., R. bicolor Urban, etc.). Either convergent evolution is oper- ating ?among these species or section Duchartrea may be more closely related to Rhytidophyllum than other sections in Gesneria. Rhytidophyllum, consisting of about 20 species, inhabits the islands of Cuba, Jamaica, Hispaniola, Puerto Rico, and Grenada in the West Indies, and in northern South America, the countries of Colom- bia and Venezuela. T h e genus can be divided into two large groups on the basis of corolla shape, par- alleling the primary division in Gesneria. Rhytido- phyllum will not be treated further here. T h e species included in the Gesnerieae are con- sidered herein as biological species in the sense of Solbrig (1970:112), ?a group of interbreeding populations reproductively isolated from any other such groups of populations.? The species may also be distinguished morphologically by a series of correlated characters from any other species. Reproductive isolation is the criterion for biological species. Species of Gesneria may hy- bridize in nature, but probably because of post- fertilization isolation mechanisms, the hybrids seldom develop and persist, allowing the species to remain distinctive in ?their ecological and evo- lutionary roles? (Grant, 1971). Mechanisms for isolation that appear to be operating in Gesneria include some of those given by Banks (1970) and Solbrig (1970): (1) geographical separation, (2) ecological separation, (3) pollen incompatibility, (4) structural differences in flowers requiring specific pollinators, (5) different flowering times, (6) gametic incompatibility, (7) hybrid sterility, (8) hybrid weakness, and (9) lack of hybrid establishment. The criterion for distinguishing subspecies within the species has been the presence of morphological variants appearing in two or more geographically distinct populations. Some species of Gesneria have been divided into varieties because the plants are less morpho- logically distinct or geographically isolated than those taxa considered subspecies in other species. Key to the Genera in Tribe Gesnerieae 1. Capsule rostrate, the rostrum curved, equaling or exceeding the length of the floral tube; leaves opposite and decussatc, bases of petioles joined across the stem; Cuba and Jamaica . . . . Pheidonocarpa, new genus 1. Capsule not beaked, if raised then the free portion not equaling or exceeding the floral tube; leaves alternate, rarely approximate, or appearing so on young or slowly growing branches, exceptionally opposite, but then petiole bases not joined across the stem. 2. Costae absent on the capsule; capsule villous, seldom verrucose or glabrous (but sometimes glabrcscent in age) and rarely with sepals over 1.2 cm long; filaments adnate to the base of the corolla tube for a distance greater than the basal thickness of the filament, except in Xhytidophyllurn cumanense; leaves areolatc, tomentose-hirsute, or rarely glabrescent, but then the corolla tubular and red 2. Costae present on the capsule, or the capsule verrucose or glabrous, but not villous unless the sepals are over 1.2 cm long (Gesneria gloxinioides); filaments adnate to the base of the corolla tube for a distance no greater than the basal thickness of the filament: leaves glabrous or scabrous to pilose, smooth, plane or rarely bullate . . . . . . . . . . . . . . . . . . . . . . . . . Gesneria ....................................................................................................................... ............................................................ Rhytidophylluin 40 SMITHSONIAN CONTRIBUTIONS T O BOTANY Pheidonocarpa L. Skog, new genus Gesneriae L. et Rhyt idophyl lo Mart. affinis, foliis oppositis decussatis et capsulis rostratis, rostro curvo tubum floralem aequante vel ex- cedente differt. Plants terrestrial, perennial, suffruticose, fibrous- rooted: stems woody, erect or decumbent. Leaves opposite, decussate: petioles short, bases of opposite leaves joined across the stem, vascular trace an ?invaginated crescent? (fide Wiehler); blades somewhat rigid, scabrous. Inflorescences axillary, each a compound di- chasium: peduncles much exceeding the subtending leaves; flowers numerous, protandrous, zygo- morphic; floral tube turbinate, pilose; calyx lobes 5, briefly connate at base; corolla tube ventricose, limb 5-lobed; stamens 4, filaments adnate to base of corolla tube, but free above, anthers coherent or free, exserted at anthesis; ovary half-inferior, disc annular, 5-lobed. Capsule turbinate, pilose, dry, splitting into two valves from the curved rostrate apex; seeds nu- merous, minute, fusiform, striated. TYPE-SPECIEs.-Pheidonocarpa corymbosa (Swartz) L. Skog. DrscussIoN.-Until now the tribe Gesnerieae has included only the two closely related genera, Ges- neria Linnaeus and Rhy t idophy l lum Martius, which are similar in having leaves arranged alter- nately, roots fibrous, ovaries inferior, chromosome numbers of usually n = 14, and being limited to the Caribbean region and northern South America. In 1788, Olof Swartz described among his new species of Gesneria a new plant from Jamaica, Gesneria corymbosa, later transferred by Urban (1901) to Heppiel la Regel. I n 1957, C. V. Morton described from Cuba another species, Heppiel la cubensis. All other species of Heppiel la are known only from South America where their apparent center of distribution is from Colombia to Ecuador. Recognizing the apparent disjunct distribution, Wiehler (1971) removed Heppiel la corymbosa and H . cubensis from the rest of Heppiel la , citing as additional reasons: (1) the lack of scaly- rhizomes as are found in the South American species of Heppiel la; (2) a chromosome number of n = 14 (Lee, 1966), unknown in the tribe Achimeneae where other species of Heppiel la reside; (3) the stomata of abaxial leaf surfaces are grouped into ?islands,? also unknown in other members of the tribe Achimeneae; and (4) the vasculature pattern of the petiole differs from other species of Heppiel la . Wiehler, however, transferred the two West Indian species to Gesneria, citing as evidence (in addition to the similar chromosome number, petiole vascular pattern, stomata1 arrangement, and fibrous roots) the occurrence of apparently opposite, decussate leaves on a single plant of a species of Gesneria in cultivation. I concur with Wiehler?s decision to exclude the West Indian Heppiel la species from the continental species. But his decision to unite the plants with Gesneria cannot be accepted, particularly on the basis of the leaf arrangement. Truly opposite and decussate leaves are not known for any species of Gesneria or Rhyt ido- phy l lum, except for the apparently aberrant form examined by Wiehler. This phenomenon is dis- cussed under ?Morphology and Anatomy.? Other characters besides leaf arrangement that distinguish species of Gesneria from Pheidono- carpa are listed in Table 1, but notably the most important are the differences in crystal types, in- florescence form, ovary position, and length of capsule beak. Because of the similarities of the West Indian species formerly in Heppiel la to the members of the tribe Gesnerieae, the concept of the tribe has been emended to include plants with opposite decussate leaves and flowers with half-inferior ovaries, but these species are retained in a separate genus within the Gesnerieae. The name Pheidonocarpa is derived from the Greek pheidon (= oilcan) 4- karpos (= fruit), alluding to the shape of the fruit. Only one species of Pheidonocarpa with two subspecies is known. 1. Pheidonocarpa corymbosa (Swartz) L. Skog, new combination Subshrubs: stems and branches to 60 cm long, 2-6 mm in diameter, bark reddish and pubescent, pilose to arachnoid when young, becoming gray and glabrescent with age, internodes 0.6-4.0 cm long, apex somewhat resinous, nodes swollen, developing interpetiolar ridges. Leaves opposite: petioles sulcate to terete, sub- NUMBER 29 sessile or to 11 mm long, 1-2 mm across, reddish to green, pubescent with glandular or eglandular 1- to many-celled hairs; blades ovate to elliptic, 3.6-8.7 cm long, 1.64.7 cm wide, base subcordate, truncate to acute, margin serrate to crenate- dentate, apex acute, adaxial surface \grayish green, scabrous from calcium-containing cells in the bases of the hairs, abaxial surface lighter green, pubescent with short dense trichomes, with some glandular trichomes on veins, veins prominent, stomata in distinct groups. Inflorescences of many flowers, in a somewhat condensed compound dichasium: peduncles te- rete, 10.8-35.0 cm long, red, pubescent; bracts at the apex of the peduncles, lanceolate, 2-4 mm long, ca 1 mm wide, green, pubescent; pedicels 0.4-1.6 cm long; floral tube turbinate, 1-2 mm long, 1-3 mm in diameter, reddish, glandular- pubescent; calyx lobes spreading, triangular, con- nate at base for less than 1 mm, 1 4 mm long, green to reddish and pubescent with glandular and eglandular trichomes outside, green and pubescent inside, veins not evident; corolla tube curved, ventricose, slightly gibbous at base, 1.4-2.1 cm long, 2-4 mm wide at base, outside dark red-orange to yellow, pubescent, glandular, inside reddish, glabrous, limb bilabiate, lobes 5, 2-7 mm long, 2.5-5.0 mm wide, reddish or yellow with darker reddish lines, glabrous except for sparse glandular hairs at mouth; stamens 4, exserted about 2 mm beyond corolla mouth, filaments to 2.2 cm long, yellow, sparsely pubescent near base, glabrescent toward apex, anthers oblong, 1.0-4.5 mm long, 1-2 mm wide, coherent or free, connective reddish; ovary apex pilose, style puberulent, red, stigma bilobed. Capsule turbinate, 7-9 mm long, 3-6 mm in diameter, reddish, becoming gray, costae visible near base, if at all. The habitats of the two subspecies are appar- ently very similar. Each is restricted to nearly perpendicular limestone cliffs of a river gorge. The E A S T E R N CUBA zoo I 200 km I J A M A I C A 41 N t k 15O FIGURE 23.-Distribution of Pheidoncarpa corymbosa in Ja- maica and Cuba. ( P . c. subsp. corymbosa = solid circles; P . c. subsp. cubensis = open circles.) primary pollinator of each subspecies is probably a hummingbird. The flowers display some of the characteristics of flowers pollinated by humming- birds with red corolla tubes oriented horizontally. A pollinator must have a bill that nearly equals or is greater than the length of the corolla tube (1.4-2.1 cm) to reach the nectar at the base of the tube. In Jamaica, possible pollinators are Mellisuga minima (Figure 25), Anthracothorax mango, and Trochilus polytmus, whose bills have a mean length of 1.2, 2.6, and 2.1 cm, respectively (from specimens measured at CU and US). The shape of the head and bill of Trochilus polytmus more closely approximates the shape and size of the corolla of P. corymbosa subsp. corymbosa, and is Key to the Subspecies of Pheidonocarpa corymbosa 1. Calyx lobes to 2 mm long, narrowly triangular; anthers 1-2 mm long, usually coherent in 2 pairs; peduncles 10.8-13.9 cm long, 1-3 times as long as the subtending leaf; Jamaica . . . . . . . . . . . ......................................................................................................... la. subsp. corymbosn mm long, broadly triangular; anthers to 4.5 mm long, usually free; peduncles 18-35 cm long, 4 or more times as long as the subtending leaf; Cuba ........................................ 1. ............................................................ . . lb . subsp. cubensis, new combination and rank 42 SMITHSONIAN CONTRIBUTIONS T O BOTANY FIGURE 24.-Representative specimen of Pheidonocarpa corym- bosa (Swartz) L. Skog, subsp. corymbosa from Jamaica (Proc- tor 10205, F). presumed the pollinator, This bird is known from the range of the plants. l a . Pheidonocarpa corymbosa subsp. corymbosa Gesneria corymbosa Swartz, Prodr. 89, 1788.-Sw., F1. Ind. Pentarhapia ?corymbosa (Swartz) Hanstein, Linnaea 34:307, Heppiella corymbosa (Swartz) Urban, Symb. Ant. 2:368, Occid. 2:1022, 1800,-DC., Prodr. 7:528, 1839. 1865. 1901.-Adams, F1. PI. Jamaica 682, 1972. Young stems and branches pubescent to pilose Leaf margin serrate. Peduncles 10.8-13.9 cm long; pedicels 0.4-1.4 cm long; calyx lobes narrowly triangular, to 2 mm long; upper lobes of the corolla limb 4-5 mm long, with whitish hairs. ca 2.5 mm wide, lateral lobes ca 3 mm long, ca 3.5 mm wide, lower lobe ca 2 mm long, 3 mm wide, margin erose to dentate; anthers 1-2 mm long, connective often dark red, usually coherent in 2 pairs, pollen grains isopolar, prolate, size small, 18.5-20.0 pm long at the polar axis, ca 8.5 pm long at the equatorial axis, tricolpate, apocolpia trun- cate to rounded, colpi narrowly attenuate, sexine uniformly heterobrochate, reticulate, lumina ca 0.25 pm across (Figure 1%). TYPE-CoLLECTIoN.-?India occid.: Jamaica,? 0. Swartz sn (S, holotype). CHROMOSOME NUMRER.-~ = 14 (Lee, 1966a). rymbosa subsp. corymbosa has been found only in the gorge of the Cane River in eastern Jamaica (Figure 23), where it grows on shaded limestone cliffs (Figure Id) from 80 to 150 m elevation. Flowering occurs throughout the year. SPECIMENS ExAMINED.-JAMAICA. PARISH OF ST. ANDREW: Cane River Gorge, ca 400 ft, 28 April 1960, C. Adams 6967 (UCWI); 100 m, 12 July 1962, F. Fosberg 42635 (US): 400 f t , 12 July 1907, IV. Harris 9621 (BM 2 sheets, C, F, K , NY 2 sheets, UCWI, US); 250 ft, 18 December 1907, W . Harris 10053 (BM 2 sheets, C, F, K, NY, UCWI, US); 100 ft, 9 No- vember 1912, W . Harris I1427 (BM, F, NY, UCWI, US); 1 January 1916, W . Harris 12314 (BM, F, GH, K, MO, NY, UCWI, US); no date, U . Morley 387 (UCWI); 14 February 1948, A . van der Porten sn (IJ): 250-400 ft, 22 May 1955, G. Proctor 10205 (BM, F, IJ, MO, US); 500 ft, February 1957, E. Robertson 2996 (K, UCTVI); 400 ft, 9 August 1970, L. Skog 1620 (BH, US); 18 December 1955, W. Stearn 3 (BM); 8 April 1956, W . Stearn 694 (A, BH, BM); 300-500 ft, 9 March 1958, T. Yuncker 18349 (BM, MO). LOCALITY UN- KNOWN: ?Gesneria aspera, Jan.,? [1821], C. Bertero sn (AWH); ?Jamaica, 1784-1786,? 0. Swartz sn ( S ) ; ?India occid.: Ja- DISTRIBUTION A D EcoLocY.-Pheidonocar~a CO- FIGURE 25.-Flower of Pheidonocarpa corymbosa subsp. corymbosa with a preserved specimen of a possible pollinator, Mellisuga minima. NUMBER 29 43 maica,? no date, 0. Swartz sn ( S , holotype of Gesneria corym- bosa Swartz). CULTIVATED: Cornell University, G-839, 31 March 1965, R. Clark sn (BH), 24 September 1965, B . Jack- son sn (BH); Cornell University, G-1367, 11 May 1972, L . Skog 1826 (BH). Drscuss1oN.-The typical subspecies has been col- lected recently from a single locality, and pre- sumably the type also came from the gorge of the Cane River, St. Andrews Parish, Jamaica. Plants have been brought into cultivation at least twice and grown at Cornell University under acces- sion numbers G-839 and G-1367. lb. Pheidonocarpa corymbosa subsp. cubensis (Morton) L. Skog, new combination and rank Heppiella cubensis Morton, Brittonia 9:21, 1957. Gesneria mortonii Wiehler, Baileya 18:4, 1971 [non Gesneria cubensis (Decaisne) Baillon (ISSS)]. Young stems pilose to arachnoid. Leaf margin crenate-dentate. Peduncles 18-35 cm long; pedicels 1.1-1.6 cm long; calyx lobes broadly triangular, to 4 mm long; corolla with upper lobes 4-7 mm long, ca 4 mm wide, margin erose, lateral lobes ca 5 mm long, 5 mm wide, margin entire, basal lobe 3 mm long, ca 5 mm wide, margin entire; anthers to 4.5 mm long, usually free. TYPE-COLLECTION.-Rio Jauco, Oriente Province, Cuba, Bro. Leon 11685 (NY, holotype (Figure 26); LS, NY, US, isotypes). DISTRIBUTION A D EcoLocY.-Pheidonocarpa co- rymbosa subsp. cubensis is known only from the Oriente Province of Cuba, where it grows on lime- stone cliffs from 50-300 m elevation. Flowering probably occurs throughout the year. SPECIMENS ExAMINED.-CUBA. PROVINCE OF ORIENTE: Loma del Esparto, Cajobabo, 31 December 1959, Alain Liogier & M . Lopez F . 7105 (US); Cueva del Toro, Caleta, east of Jauco, 200-300 m, 17 J u l y 4 August 1924, Bro. Leon 11812 (NY 2 sheets, US); rocky banks of Jauco River, Jauco, 17 J u l y 4 August 1924, Bro. Leon 11685 (NY, holotype of Heppiella cubensis Morton; NY, US, isotypes); dry cliffs, Jauco, May 1940, Bros. Leon & M . Victorin 17748 (US); dry cliffs beside the Rfo Jauco, ca 50 m, 15-16 January 1956, C. Morton & Alain Liogier 9154 (US). DIscussroN.-This taxon from Cuba (Figure 23) is included here as a subspecies of Pheidonocarpa corymbosa because of the similarity in habitat and morphology to the typical subspecies. FIGURE 26.-Holotype of Pheidonocarpa corym bosa Heppiella cubensis Morton and subsp. cubensis (Morton) I,. Skog, new combination and rank, Leon 11685 (NY). Gesneria Linnaeus Gesneria Linnaeus, Gen. P1. ed. 5, 267, 1754.-Linn. Sp. P1. ed. 1, 612, 1753. Pentarhaphia Lindley, Bot. Reg. 13: sub. pl. 1110, 1827 [(?Pentaraphia?), non Pentarrhaphis Humboldt, Bonpland & Kunth (1816)l. [Type-species: Pentarhafihia longifloru Lindley (=Gesneria ventricosa Swartz).] Conradia Martius, Nov. Gen. & Sp. 3:38, 1829 [nom. illeg., non Rafinesque (1825), nec Nuttall (1834), nec Kufferath (1914)l. [Type-species: Pentarhaphia longiflora Lindley (= Gesneria ventricosa Swartz), vide Morton & Denham (1972).] Duchartrea Decaisne, Ann. Sci. Nat. Bot. ser. 3, 6:109, 1846. [Type-species: Duchartrea viridiflora Decaisne.] Herincquia Decaisne ex Herincq, Rev. Hort. ser. 3, 2:322, 1848. [Type-species: Herincquia floribunda (Lemaire) De- caisne ex Herincq (= Gesneria libanensis Linden ex Mor- ren).] 44 SMITHSONIAN CONTRIBUTIONS TO BOTANY Ophianthe Hanstein, Linnaea 26:205, 1854. [Type-species: Ophianthe libanensis (Linden ex Morren) Hanstein.] Chorisanthera (G. Don) Orsted, Cent. Gesn. 35, 1858. [Type- species: Chorisanthera pumila (Swartz) Orsted (=Gesneria pumila Swartz).] Codonoraphia Orsted, Cent. Gesn. 68, 1858. [Lectotype-species: Codonoraphia calycosa (W. Hooker) Orsted.] Vaupellia Grisebach, F1. Brit, W. Ind. 460, 1862. [Type- species: Vaupellia calycina (Swartz) Grisebach.] Plants fibrous-rooted, perennial, herbaceous, suffruticose with decumbent, pendent or erect stems and branches, fruticose with erect stems or with spreading basal branches to arborescent with stems to 10 m tall: bark glabrous to pilose, resin often persistent on stem, leaves and inflorescences, pith green to red, cells containing crystals; branches with internodes cylindrical or rarely flattened, nodes usually swollen, unilacunar with one trace. Leaves dorsiventral, a1 ternate, rarely approxi- mate, opposite and decussate in one population of Gesneria fruticosa, spirally arranged in a 215 phyllotaxy, often crowded at branch apices: pet- ioles, if present, terete to sulcate, green to reddish- brown, glabrous to pilose, occasionally verrucose, resinous, vascular trace a cylinder or U-shaped in transverse section; blades ovate or lanceolate to obovate, oblanceolate or suborbiculate, membra- nous to coriaceous, plane to bullate, base cuneate to truncate or cordate, margin entire, crenate, dentate, serrate, or lobulate, occasionally ciliate, apex retuse to caudate, adaxial surface usually dark green, glossy or dull, abaxial surface lighter green to reddish-brown, both sides glabrous to pilose or scabrous, trichomes, when present, uni- seriate, glandular or nonglandular, veins pinnate, prominent at least below, arcuate at margin, epi- dermal cells with shallow or nonundulating walls, stomata not raised, cruciferous-type of 2 guard cells and 3 subsidiary cells, internally an adaxial multi- ple epidermis usually present with crystal con- taining cells. Inflorescences of many flowers in a compound dichasium or in some species reduced to a single flower, shorter than or exceeding the subtending leaves: peduncles usually terete, rarely quad- rangular, green to red, glabrous to pilose, occa- sionally verrucose, resinous; bracts 2, fertile or sterile, leaflike or reduced; pedicels similar to the peduncles, green to red, glabrous to pilose, usu- ally resinous; flowers protandrous or protogynous; floral tube obconic, turbinate, or nearly spherical, green to reddish-brown, glabrous to villous, occa- sionally verrucose, resinous; calyx lobes 5 or rarely 4 or 6, distinct or briefly connate above the ovary, each lobe erect to patent, sulcate or flat, ovate to lanceolate or filiform, white, green to red or brown, glabrous, or puberulent to pilose, margin usually entire, seldom toothed; corolla tubular, cylindrical, ventricose, or infundibuliform- campanulate, white, green, red, orange, yellow to brown, glabrous to pilose, sometimes resinous and verrucose, limb 5-lobed, occasionally two-lipped, each lobe erect to reflexed; fertile stamens usually 4, with an infertile staminode, rarely 5 stamens fertile, filaments of equal length, linear, curved to geniculate at the base, adnate to corolla tube for a distance no more than the basal thickness of the filament, white, green, yellow or red, glabrous or pilose below, connective often prolonged beyond anther, anthers sagittate to oblong, usually co- herent at anthesis in two pairs at their apices or in a row side by side, becoming free, 2-celled, opening by longitudinal slits, pollen isopolar, prolate, size small to medium, sexine usually reticulate, seldom smooth or verrucose; pistil 1, hemiparacarpous, ovary inferior or rarely subinferior, 2-carpelled, unilocular, apex pubescent or glabrous, ovules numerous, anatropous, integument single, disc annular or undulating surrounding the stylar base, style usually as long as the stamells or slightly longer, curved, white, green, yellow or red, gla- brous to pilose, stigma stomatomorphic, clavate, capitate, or bilabiate, usually papillate. Capsule loculicidal, turbinate or nearly spheri- cal, dehiscent from the apex into 2 or 4 valves, or forming a splash cup, reddish-brown to gray, glabrescent, costae 5-15 or capsule verrucose and costae obscure; seeds numerous, linear, or fusi- form, to rhombic or oblong, striate, twisted, tawny, red, brown or black; cotyledons 2, equal. LECToTYPE-SPECIEs,-GeSneria humilis Linnaeus, vide Fritsch (1894), Morton & Denham (1972). Gesneria is here divided into nine sections, dis- tinguished by factors related to reproduction, i.e., characters of flowers and fruit, supplemented by correlated vegetative characters. The sections appear to be natural groups of species sharing many characters, but also clearly related to other sections or species within Gesneria. Such relation- NUMBER 29 Key to the Sections of Gesneria 45 1. Diameter of the corolla mouth less than or equal to the diameter of the middle of the tube, the tube nearly cylindrical or ventricose, or the tube narrowly funnelform expanding gradually from the base (section Stenochonanthe), or hypocrateriform (section Pentarhaphia), reddish or occasionally yellow to green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Plants erect, shrubs or small trees, usually more than 60 cm tall ...................................... 3 3. Corolla verrucose: stem unbranched, or branched only at the base; calyx and capsule verruculose to verrucose; Cuba . . . . . . . . . . . . . . . . . . . . . . . Section 4 . Myrmekianthe, new section 3 . Corolla not verrucose; stem usually much branched; calyx and capsule smooth, seldom verruculose ............................................................................................................................... 4 4. Leaves lobulate; axillary buds and young branches lanate, not resinous . . . . . . . . . . . . . . . . . . .................................................................................. Section 3. Lachnoblaste, new section 4 . Leaves entire to dentate-serrate; buds and young branches glabrous to pilose, usually resinous .............................................................................................................. 5 5 . Stamens included; corolla evenly expanded from the base; Jamaica .................... .................................................................... Section 2. Stenochonmthe, new section 5 . Stamens exserted and corolla ventricosc or widest at the base or middle, or if stamens included, then corolla not expanded, but hypocrateriform (Gesneria aspera, G. haitiensis), or corolla suburceolate (G. pulverulenta) ......................... 2. Plants nearly stemless, often decumbent or pendent, rarely to 60 cm tall .......................... G 6 . Inflorescences projecting beyond the subtending leaves; corolla yellow, orange, or green to white, red only in Gesneria brachysepala; leaves plane . . . . . . . . . . . . . . . . Section 5 , Gesneria 6 . Inflorescences shorter than the subtending leaves; corolla usually red, seldom orange- red, rarely yellow; leaves bullatc, rarely nearly plane ......................................................... 1. Diameter of the corolla mouth larger than the diameter of the middle; the tube infundi- buliform-campanulate, green to yellow or brown, rarely reddish to purple ..................... . 7 7 . Plants erect, shrubs or small trees well over 60 cm tall; inflorescences exceeding the sub- tending leaves, of one to many flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8. Stem unbranched, unless at the base, verrucose and nearly glabrous above; stamens 8. Stem usually much-branched and smooth, sometimes pubescent, if glabrous, then obscurely if at all verruculose; stamens seldom included; leaves membranous to sub- 7 . Plants nearly stemless or decumbent, rarely up to 60 cm tall; inflorescences shorter than the subtending leaves, of one to four flowers . . . . . . . Section 9. Chorisanthera, new transfer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Section 1. Pentarhaphia ....................................................................................... Section 6. Physcophyllon, new section included; leaves coriaceous ................................................................... Section 8. Duchartren coriaceous .................................................................... Section 7 . Dittanthera, new transfer ships can be seen between sections Duchartrea and Myrmekianthe, Chorisanthera and Physcophyllon, and Pentarhaphia and Dittanthera. The genus may be divided into two large groups on the basis of corolla shape. The larger group, consisting of six sections, has tubular and nearly cylindrical or ventricose corollas. This type of corolla is believed to be adapted for hummingbird- pollination. The smaller and perhaps more ad- vanced group, consisting of three sections, has campanulate corollas, abundant pollen, and nectar possibly attractive to bats. However, the bats have not yet been actually observed pollinating. Evolution in the genus may have occurred in a series of reductions. An example of a more primi- tive species is Gesneria ventricosa, a species of shrubs with red, tubular, hummingbird-pollinated flowers. From such an example two major lines of evolution appear to have developed: (1) shrubs or trees with hummingbird-pollinated flowers, (2) acaulescent or short-stemmed herbs or suhshrubs with hummingbird-pollinated flowers. Both of these lines may have given rise to bat-pollinated species. Relationships among the sections will be dis- cussed under the sections concerned, together with the relationships among the species in each section. Gesneria, when described by Linnaeus in 1753 and 1754, was composed of two species, G. humilis and G. tomentosa. Neither species was chosen as the type-species at that time, Gesneria tornentosa was transferred to Rhytidophyllum by Martius in SMITHSONIAN CONTRIBUTIONS T O BOTANY 1829 and selected as the lectotype species for that genus by Britton and Wilson (1925). Fritsch (1894) was apparently the first to retain G. humilis in Gesneria and refer G. tomentosa to Rhytido- phyllum, thus Gesneria humilis Linnaeus has be- come the residual lectotype species for Gesneria (Morton and Denham, 1972). Conradia Martius was an intentional, but un- justified, renaming of Pentarhaphia Lindley (Mor- ton and Denham, 1972). According to Article 7 of ICBN a nomenclaturally superfluous name ?is automatically typified by the type of the name which ought to have been adopted under the rules? (ICBN, 1972: 19). Thus, the only species and type of Pentarhaphia when it was published, P. longiflora Lindley (which itself is a superfluous name for Gesneria ventricosa Swartz), became the type of Conradia rather than Conradia humilis, selected by later authors. Until the present time, Codonoraphia Orsted has not been typified. T o stabilize the name, Codonoraphia calycosa (W. Hooker) Ozhhhvrsted has been selected as the lectotype species. Section 1. Pentarhaphia (Lindley) Fritsch Pentarhaphia (Lindley) Fritsch in Engl. & Prantl, Nat. Pflanzenfam. 4(3b):184, 1894. Pentarhaphia section Eupentarhaphia Bentham in Bentham & Hooker, Gen. P1. 2:1005, 1876. [Type-species: Pentar- haphia longiflora Lindley (= Gesneria ventricosa (Swartz).] Gesneria section Conradia (Martius) Fritsch in Engl. & Prantl, Nat. Pflanzenfam. 4(3b):184, 1894 [as to type, not as to description]. Small subshrubs to large shrubs: stems erect, to 4 m tall, resinous: branches erect, or often divari- cate arising from base or from upper leaf axils. Leaves lanceolatc, elliptic, ovate to oblanceolate or obovate, occasionally falcate, submembranous to coriaceous, plane, glossy or dull, margin some- times recurved. Inflorescence 1 to 4 (-many)-flowered, shorter than, equaling or exceeding the subtending leaves, occasionally verrucose: corollas tubular, ventricose at the middle and narrowing to mouth or sub- urceolate or hypocrateriform, yellow, orange to red; stamens usually exserted, seldom included. Capsules turbinate to nearly spherical, smooth, costae usually prominent, sometimes alate. TYPE-SPEcIEs.-Pentarhaphia longiflora Lindley (= Gesneria ventricosa Swartz). DIscussIoN.-Section Pentarhaphia comprises a large complex of species which share many similar characters. It resembles and is probably closely re- lated to section Stenochonanthe. The floral struc- tures of the plants in the two sections are very similar, as are the vegetative characters. This simi- lar development may, however, be the result of parallel evolution and response to hummingbird pollination. A resemblance in corolla form can also be seen in some species of Rhytidophyllum (e.g., R. berteroanum Martius), but these differ in other essential characters. Ancestors of this section may have been the stock from which many of the sec- tions evolved, but the Pentarhaphia line has main- tained seemingly primitive characters. Members of this section are mainly concentrated on Hispaniola with a few species in Cuba, Jamaica, and the Lesser Antilles (Figures 27, 29, 36, 43, and 44). Key to the Species of Section Pentarhaphia 1. Abaxial leaf surfaces, petioles, peduncles, and corollas clothed with pilose or villous trichomes, and resinous. 2. Peduncles more than 1.8 cm long; floral tube turbinate. 3. Corolla red, limb patent; adaxial leaf surfaces asperate; inflorescences of one flower; Haiti .................................................................................................................................. 2. G. aspern 3. Corolla yellow, limb erect or porrect; adaxial leaf surfaces glabrescent: inflorescences 2. Peduncles 1 cm or less long; floral tube obconic: Dominican Republic .. 15. G. paruifolia 1. Abaxial leaf surfaces, petioles, peduncles, and corollas glabrous, or scarcely pubescent near petiole or peduncle base, or in petiole sulcus, but resinous. 4. Leaves subtending the inflorescences more than twice as long as the inflorescence. 5 . Leaf margins entire, blades 3.6 cm long or less; Cuba: Province of Piaar del Rio ........ 5. Leaf margins denticulate, serrulate, crenulate, dentate, crenate, or serrate, blades of three to several flowers; Cuba ..................................................................... 7 . G. glandulosn ....................................................................................................... usually much more than 4 cm long, largest leaves more than 6 cm long. NUMBER 29 47 6. Bark subrugose, verrucose, verruculose, or tuberculate, lenticels obvious; inflores- cences usually of one flower, if more then leaf margins crenulate. 7. 9. G. hmrisii 7 . Style pilose; leaf margin crenulate or denticulate; Cuba. 8. Floral tube globose, costae 10; leaves obovate, 4.4-10.0 cm long; Cuba: Province of Las Villas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. G. clarensis 8. Floral tube narrowly obconic, costae 5; leaves lanceolate, elliptic, or seldom Style glabrous; leaf margin crenate or dentate above middle; Jamaica . . . . . . . . . . . . . . ....................................................................................................................... obovate, 2.0-6.8 cm long; Cuba: Province of Oriente . . . . . . . . . 10. G. heterochroa 6. Bark smooth, lenticels obscure; inflorescences of two or three flowers. 9. Leaves narrowly elliptic to oblanceolate, 1.5-3.3 cm wide, denticulate above the . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13. G. lanceolatn 9. Leaves broadly elliptic to obovate, 3.1-7.0 cm wide, serrate above the middle; Haiti: Dkpartement du Sud . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..14. G. odontophylla 4. Leaves subtending the inflorescences less than twice as long as the inflorescences to much shorter. 10, Corolla tube yellow or greenish-white externally above the base; stamens exserted only 9. G. horrisii 12. Capsule turbinate, ca 4 mm long; leaves 2.4-4.2 cm long, serrulate or denticulate above the middle, base narrowly acute; branches few; Haiti ..11. G. hypoclada 12. Capsule narrowly obconic, to 8 mm long; leaves 2.5-11.2 cm long, margin crenate or serrate above, base obtuse, branches numerous; Cuba ....................................... 10. Corolla tube red, red-orange or rose-pink externally, if yellow, then only at the base. 13. Corolla limb patent; inflorescences of one flower, peduncles very short with pedicels more than twice as long; Haiti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8. G . haitiensis 13. Corolla limb with some lobes erect, reflexed, porrect, or spreading, but not all patent; inflorescences of one to many flowers, peduncles usually exceeding length of pedicels, if pedicels lacking, peduncles more than 1 cm long. 14. Peduncles and pedicels together 1.3 cm long or less, not arcuate, and stamens middle; Haiti: Dkpartement de l'Ouest to limit of corolla tube. 11. Style glabrous; bracts lacking; Jamaica . . . . . . . . . . . . . . . . . . . . . . . . 11. Style pilose; bracts present, but sometimes caducous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..17c. G. salicifolia var. ferruginea exserted 3 mm or less; corolla tube suburceolate; Dominican Republic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. G. pulverulenta 14.Peduncles and pedicels together usually more than 2 cm long, often much longer, and peduncles usually arcuate, if shorter or not arcuate, then stamens exsertcd more than 4 mm at anthesis; corolla tube usually ventricose. 15. Stamens exserted beyond corolla limb; peduncles arcuate or not. 16. Ovary apex glabrous; corolla limb lobes erect, tube maculate inside; Cuba: Province of Oriente .................................................................. 3 . G . binghamii 16. Ovary apex tomentose to pilose; corolla limb erect to reflexed, rarely all lobes erect, tube not maculate. 17. Peduncles 2.0-15.9 cm long; leaves with petioles 0.5-4.5 cm long, blades 4.1-22.7 cm long; inflorescences of one to many flowers, usually two to four or many, seldom only one; bracts 0.6-1.5 cm long; Lesser Antilles . . .......................................................................................... 18. G. ventricosa 17. Peduncles 0.4-2.0 cm long; leaves with petioles 0.2-1.2 cm long, blades 1.1-7.0 cm long; inflorescences of usually one, rarely two or three flowers: bracts 0.1-0.5 cm long; Cuba, Hispaniola . . . . . . . . . . . . . 6. G . cubensis 15. Stamens included or exserted only to limit of corolla limb; peduncles arcuate. 18. Corolla 1.0 cm long or less, ventricose at base, narrowing toward middle; inflorescences of one or rarely two flowers; Jamaica ....... 12. G . jamaicensis 18. Corolla 1.1 cm long or more, ventricose at or near the middle; inflores- cences of usually more than one flower. 19. Leaves entire toward the base, denticulate or serrulate toward apex; corolla yellow internally; Cuba: Provinces of Oriente and Pinar del Rinar ................................................................................. 17. G. salicifoliu 19. Leaves sharply serrate; corolla red internally; Cuba: Province of Oriente .................................................................................. 19. G. wrightii SMITHSONIAN CONTRIBUTIONS T O BOTANY H I S PA N I0 L A FIGURE 27.-Distribution of Gesneria section Pentarhaphia in Hispaniola. (G. aspera = open triangles; G. cubensis var. cubensis = solid circles; G. cubensis var. truncata = solid squares; G. hi t iens is = solid triangles; G. hypoclada = open stars within circles: G. lanceolata = open squares; G. odontophylla = solid stars; G. paruifolia = open stars; G. pulverulenta = open circles.) 2. Gesneria aspera Urban & Ekman Gesneria aspera Urban & Ekman in Urban, Ark. Bot. 20A(5): 49, 1926. Shrubs or small trees: stems woody, erect, bark when young reddish and invested with numerous reddish, few-celled patent trichomes, becoming gray-brown and glabrous, resinous at apex, inter- nodes 0.1-1.9 cm long; branches many, 3 mm in diameter at 10 cm below apex, lenticels erumpent, nodes slightly swollen. Leaves approximate to alternate: petioles terete near node, becoming flattened near base of blade, 1-4 mm long, ca 1 mm wide, reddish and covered by the patent glandular hairs; blades ovate to obovate, 1.5-2.4 cm long, 0.8-1.4 cm wide, stiff and coriaceous, base rounded to obtuse, margin entire near the base, becoming crenate-dentate toward the acute apex, adaxial surface bullate, light green, asperate from numerous stiff bulbous- based white hairs, lateral veins 3 to 5, slightly immersed, abaxial surface tawny except for the reddish prominent veins, pilose. Inflorescences axillary, I-flowered, about twice as long as the subtending leaves: peduncles 1.8-3.0 cm long, 0.3-0.4 mm in diameter, purplish-brown, pilose; bracts 2, at the base of the peduncle when young, caducous, linear, to 2 mm long, reddish, pilose; floral tube turbinate, ca 2 mm long, ca 2 mm in diameter at apex, red and pilose; calyx erect, lobes 5, barely connate at base, linear- lanceolate, 2-6 mm long, ca 1 mm wide at base, dark red, shortly and densely pilose; corolla tube hypocrateriform, 1.1-2.2 cm long, ca 0.3 cm wide at middle, outside red except orange-yellow near base, shortly and patently pilose, inside yellow with reddish lines, glabrous with prominent veins, limb patent, 7-9 mm broad, lobes orbicular, ca 5 mm long, ca 4 mm wide, upper lobes broader and slightly bifid, margin erose, glandular; stamens 4, adnate to the base of the corolla, filaments ca 15 mm long, yellowish, glabrous, anthers oblong, ca 1 mm long, coherent until anthesis, pollen yellow- white, staminode 4 mm long, lacking anther; ovary inferior, becoming half-inferior following fertiliza- tion, reddish, pilose at apex, disc annular, swollen, style ca 10 mm long, yellow, shortly pubescent, stigma clavate. Capsule shortly turbinate, extending ca 2.5 mm above calyx attachment, 5-6 nim wide, becoming gray on drying, costae 5, slightly prominent; seeds fusiform, striated, 0.6-0.7 mm long, 0.1-0.2 mm wide, brownish. TYPE-COLLECTloN.-Massif de la Selle, Morne NUMBER 29 49 Cabaio, Haiti, E. Ekman H1550 (S, holotype, Figure 28a; EHH, G, IJ, K, NY, S, US, isotypes). DISTRIBUTION A D EcoLocY.-Gesneria aspera has been found only in southeastern Haiti in the la Selle mountains on Morne Cabaio, where it grows in fissures in limestone cliffs at 1900-2000 m (Figure 27). Flowering specimens have been col- lected only in August, SPECIMENS EXAMINED: HAITI. DEPARTEMENT DE L?OUEST: Massif de la Selle, Morne Cabaio, 1900-2000 m, 20 August 1924, E. Ekman HI550 ( S , holotype of Gesneria aspera Urban & Ekman; EHH, G, IJ, K, NY, S, US 2 sheets, isotypes). DIscuss1oN.-Gesneria aspera is probably most closely allied to G. parvifolia from the Dominican Republic, but differs from it by the asperate leaves, one-flowered inflorescences, densely pilose investi- ture and the wider patent corolla limb (resembling in the last character the corollas in Gesneria reticdata and G. haitiensis). This species is known only from a single collection, perhaps be- cause of the inaccessibility of the locality. T o reach Morne Cabaio from the nearest road apparently requires ten hours travel on foot. 3. Gesneria binghamii Morton Gesneria binghamii Morton, Brittonia 9: 19, 1957.-Morton in Pentarhaphia incurva Grisebach, Cat. PI. Cub. 199, 1866. Alain & Leon, F1. de Cuba 4:455, 1957. FIGURE 28.-Type specimens: a, holotype of Gesneria aspera Urban, Ekman HI550 ( S ) ; b, holotype of Pentarhaphia incuma Grisebach and Gesneria binghamii Morton, Wright 3073 (GOET). 50 SMITHSONIAN CONTRIBUTIONS T O BOTANY Gesneria incurua (Grisebach) Urban, Symb. Ant. 2:374, 1901 [non Bentham (1845)l. Shrubs or small trees: stems woody, erect, up to 6 m tall, slender, 2-3 mm in diameter at 10 cm below apex, bark smooth, dark red and nitid above, gray below, glabrous, resinous, lenticels elongated; branches scattered, internodes 0.8-1.8 cm long, leaf scars obvious. Leaves alternate, scattered, not congested at branch apices: petioles sulcate, 3-10 mm long, ca 1 mm broad, reddish-brown, glabrous, glandular- resinous; blades lanceolate or elliptic, 3.6-9.9 cm long, 1.0-2.1(-4) cm wide, subcoriaceous, base acute, margin sharply dentate or serrate, apex acute or acuminate, adaxial surface smooth, dark green, glabrous, resinous along the midvein, abaxial surface smooth, green to brown, glabrous, resinous, veins prominent, reddish. Inflorescences of single erect flowers, about 1 1/2 times the length of the subtending leaves: peduncles arcuate, terete below, subalate above, 2.9-4.8 (-9.6) cm long including pedicels, ca 1 mm in diameter, dark red, verruculose, resinous; bracts often lacking, when present, vestigial, ca 2 mm long, ca 0.25 mm wide, red, resinous; floral tube obconic, subalate, 2-3 mm long, 2.5-4 mm wide, dark red, verruculose, nitid, resinous; calyx 5-lobed, erect but becoming incurved with age, lobes con- nate 1-2 mm at base, lanceolate, 1.0-2.1 cm long, 1.5-3 mm wide at base, outside keeled, scarlet, glabrous, resinous, inside sulcate, scarlet, glabrous, resinous; corolla tube curved, ventricose at middle, 1.4-1.8(-2.6, fide Morton) cm long, 4-5 mm wide at base, 4-8 mm wide at middle, narrowing to ca 6 mm at throat, outside bright red-orange, gla- brous, resinous, inside yellow-orange, maculate, glabrous, limb erect, 5-lobed, scarlet, margin and veins dark red, upper lobes oblong, 2-4 mm long, 2-3 mm wide, erose, glandular, lateral and basal lobes semiorbiculate, 2.0-2.5 mm long and wide, entire, glandular, mouth oblique; stamens 4, ad- nate to base of corolla for ca 0.5 mm, exserted ca 10 mm, filaments curved, ca 2.5 cm long, 1 mni wide at base, narrower above, reddish, sparsely glandular, anthers oblong ca 1.5 mm long, ca 0.75 mm wide, coherent in 2 pairs, staminode about 5 mm long; ovary inferior, apex glabrous, disc an- nular, yellow, style curved, equaling stamens in length, reddish, sparsely glandular, resinous, stigma capitate, papillate. Capsule turbinate, 6-8 mm long, 5-8 mm wide at apex, dark red to gray, glabrous, resinous, costae 5; seeds fusiform, striate, twisted, ca 0.75 mm long, ca 0.25 mm wide, black. TYPE-COLLECTION.-San Juan de Euenavista, Cuba, C. Wright 3073 (GOET, holotype (Figure 28b); BM, G, GH, K, MO, isotypes). DISTRIBUTION AND EcoLocY.-Gesneria binghamii is known from the mountains of southern Oriente Province in Cuba, where it grows on rocks in woods near rivers at 300-700 m elevation (Figure 29). Flowering material of Gesneria binghamii has been collected in March, April, August, and October. SPECIMENS EXAXIINFD: CUBA. PROVINCE OF ORIENTE: Sierra Maestra, no date, G. Bucher I0529 (NY); Sierra Maestra, headwatcrs of Rio Manacas, San Fclipe (San Juan de Ruena Vista), SW of Bueycito, 1 August 1922, E . Ekman 14705 (US), 14705b (NY), 1 4 7 0 6 ~ ( S ) ; San Juan de Buenavista, 1860-64, C. Wright 3073 (GOET, holotype of Pentarhaphia incurva Grisebach; BM, G 2 sheets, GH, K , MO, P, isotypes); Fca. ?Yao? a1 Sur de Bayamo, April 1950, C. Bingham 7247 ( I J S ) ; Rio Yao, 31 March 1943, Bro. Marie Victorin 21339 ( G H , LS); northern spur of Sierra Maestra, liest of Rio Yao, 300-700 m, 2 4 3 0 October 1941, C. Morton & J . Acuna 3492 (BM, F, G, GH, MO, NY, S, U, US 2 sheets): Sierra Maestra, foothills on north side of La Bayamesa, ca 5 km south of San Pablo de Yao, 300 m, 16 August 1951, G. Webster 4079 (GH). LOCALITY UNKNOWN: Cuba, no date, R. de la Sagra 907 (P). 4. Gesneria breuifolia Urban Gesneria breuifolia Urban, Repert. Spec. Nov. Regni Veg. 21:69, 1925. Shrubs: stems woody, erect; bark rugose, gla- brous, resinous-granulate, internodes 1-3 mm long; branches numerous, leaf scars prominent. Leaves alternate, clustered at branch apices: petioles sulcate, 3-8 mm long, ca 1 mm wide, gla- brous, resinous, blades elliptic to spathulate, 1.9-3.6 cm long, 1.5-2.3 cm wide, coriaceous, base acute, margin entire, slightly recurved, apex rotund, rarely acute or emarginate, adaxial surface plane, yellow-green to reddish-brown, glabrous, midvcin impressed, abaxial surface reddish-brown, glabrous, resinous, veins darker, prominent. Inflorescences in upper axils 1 (-3)-flowered, about one-half the length of the subtending leaves: peduncles terete, 2-5 rnm long; bracts 2, caducous, ca 1 mm long; pedicels terete, broader at apex, NUMBER 29 51 C U B A I FIGURE 29.-Distribution of part of Gesneria section Pentarhaphia in Cuba. (G. binghamii = solid squares; G. brevifolia = stars within circles; C. clarensis = open triangles; G. glandu- losa = open stars; G. heterochroa = open squares; G. salicifolia var. ferruginea = solid triangles; G. salicifolia var. salicifolia = opcn circles; G. salicifolia var. spathulata = solid circles; G. wrightii = solid stars.) 2-7 mm long, elongating to 13 mm in fruit. Flow- ers not seen. Capsule turbinate or subglobose, 4-5 mm long, 3.5-4.0 mm wide, persistent calyx lobes linear from a broad base, 5-6 mm long, glabrous, costae 5; seeds not seen. TYPE-COLLECTION.-Pan de Guajaibon, Cuba, E . Ekman 12760 (S, holotype, Figure 30; S, isotype). DISTRIBUTION A N D EcoLocY.-Endemic to the Province of Pinar del Rio in Cuba growing in dense thickets in the mountains at elevations from 750 to 800 m (fide Ekman, Figure 29). This species has only been collected in January, but in fruit. SPECIMENS EXAMINED: CUBA. PINAR DEL Rio: Pan de Gua- jaibon, on the eastern top of the mountains, ca 750-800 m, 9 January 1921, E. Ekman 12760 ( S , holotype of Gesneria brevifolia Urban: S, isotype). D~scuss~o~.-Al though this species is known only from two specimens from one locality and de- scribed without flowers, it is undoubtedly a Ges- neria, and resembles in habit and foliage other members of this section. The habit and spathulate leaves are similar to G. clarensis, whose leaves and fruits are much larger. The fruits resemble those of G. cubensis from Oriente Province and Hispan- iola (up to 7 mm long and 6 mm wide) but are smaller (4-5 mm long, 3.5-4.0 mm wide). 5. Gesneria clarensis Britton & Wilson Gesneria clarensis N. Britton & P. Wilson, Mem. Torrey Bot. Club 16:109, 1920. Tall shrubs or large trees: stems woody, bark verruculose, becoming reddish brown and resinous at apex, brown to gray and smooth below; branches 3-4 mm in diameter at 10 cm below apex, internodes 5-8 mm long, leaf scars prominent. Leaves clustered at branch apices, alternate: petioles sulcate, 0.6-1.5 cm long, 1-2 mm wide, green or reddish-brown, glandular-resinous; blades obovate, 4.4-10.0 cm long, 2.2-4.3 cm wide, sub- coriaceous, base acute, margin crenulate, apex obtuse or acute, adaxial surface smooth or rough when dry, green or reddish-brown, glabrous, veins impressed, abaxial surface reddish-brown, glandular-resinous, veins prominent. Inflorescences from upper axils, 1 to 3-flowered, less than half the length of the subtending leaves: peduncles 1.3 cm long, ca 1 mm in diameter, terete below, subalate above, reddish, glandular-resinous; bracts 2, lanceolate, 2-3 mm long, reddish, glandular-resinous; pedicels 3-7 mm long; floral tube globose, 3-4 mm long, 3-4 mm wide, reddish- brown to yellow, glandular-resinous, 10-costate; calyx 5-lobed, erect, lobes connate at base for ca 1 mm, each linear-lanceolate, 6-8 mm long, 1-2 mm wide at base, yellowish from resin, glandular on both sides, outside midvein prominent, inside sulcate; incomplete corolla tube cylindric, slightly swollen at base, broader above, length over 1 cm long, 2-3 mm wide at base, wider above, outside and inside greenish-yellow(?) glandular-resinous, limb not seen; stamens 4, adnate to base of corolla 52 SMITHSONIAN CONTRIBUTIONS T O BOTANY FIGURE 30.-Holotype of Gesneria brevifolia Urban, Ekman 12760 (S). tube, filaments curved, at least 1.5 cm long, yellowish, glabrous, anthers not seen; ovary in- ferior, apex pilose, disc annular, style curved, 1.8 cm long, reddish, pilose, stigma capitate, papillate. Capsule broadly turbinate, ca 5 mm long, ca 5 mm wide, brown, costae 5 prominent and 5 ob- scure; seeds fusiform, striate, twisted, ca 0.75 mm long, ca 0.2 mm wide, brown. TYPE-CoLLECTIoN,-~ancti-~piritus Mts, Cuba, Bros. Leon & Clemente 6504 (NY, holotype, Fig- ure 31a; NY, s, US, isotypes). DISTRIBUTION AND EcoLow.-Gesnerz'a clarensis is known only from the Province of Las Villas in central Cuba, where i t has been collected from rocky areas at 850-900 m elevation (Figure 29). The only flowering specimen was collected in January. LOCAL NAME.--"Canelon" (fide Leon & Roca 7992). SPECIMENS ExAMIKED.-CUBA. PROVINCE OF LAS VILLAS: Lomas del Banao, El Purial, on Rio Banao, ca 8.50 m, 27 January 1923, E. Ekman 16231 ( S ) ; Banao Hills, August 1918, Bro. Leon 4426 (US); Sierra del Caballete, Sancti-Spiritus Mts., 1-11 August 1916, Bros. Leon & CZemente 6504 (NY, holotype of Gesneria clarensis N. Britton & P. Wilson; NY, S, US, isotypes); Loma de la Gloria, Banao Mts., 900 m, Bro. Leon & M . Roca 7992 ([30 July 19181 NY, [31 July 19181 P); Lomas de Banao, January 1920, A . Luna 116 (NY). DIscussIoN.-The description of the available flower parts was based on Luna 116 (NY), the only specimen bearing flowers. The type, Leon & Clemente 6504, has only fruits, T h e color of the corolla is probably yellow-green, although it may be reddish since the style is definitely red. Stamens and style in other species of Gesneria are often of the same color as the corolla. Vegetatively, G. clarensis resembles G. odonto- phylla, another species described without mature flowers. The leaves are of similar size and shape, and the inflorescence is %flowered. G. odonto- phylla from the southwestern peninsula of His- paniola differs in being a small shrub with the young branches slender and divaricate, the leaf margin serrate, and the petioles to 3 cm long. 6. Gesneria cubensis (Decaisne) Baillon Gesneria cubensis (Decaisne) Baillon, Hist. Plant. 10:60, pls. 48, 49, 1888. Shrubs or small trees: stems woody, erect to 5 m tall, 2-4 mm in diameter at 10 cm below apex, bark smooth or slightly rugose, usually verrucose or tuberculate, brown to gray, glabrous, resinous toward apex, pulverulent below, lenticels elon- gated, prominent; branches mostly erect or divari- catc, reddish or brown near apex, gray or brown below, internodes 0.1-12.3 cm long, leaf scars prominent. Leaves well spaced or often conferted at branch apices, alternatc: petioles sulcate on upper surface, 2-12 mm long, 0.5-1.5 mm wide, green to reddish- brown or yellow from pulverulence, usually ver- rucose, occasionally with glandular trichomes con- gested near the base; blades elliptic to obovate or oblanceolate, 1.1-7.0 cm long, 0.5-2.7 cm wide, NUMBER 29 53 FIGURE 31.-Type specimens: a, holotype of Gesneria clarensis Britton & Wilson, Leon & Clemente 6504 (NY); b, holotype of G . truncata Alain and G . cubensis var. truncata (Alain) L. Skog, new rank, Howard 12283 (NY). somewhat rigid to coriaceous, base cuneate to acute, margin entire toward the base, crenate, serrate or dentate above, occasionally recurved, apex obtuse to acuminate, adaxial surface plane, dark green or light or reddish-brown in young leaves, glabrous, nitid, rarely foveolate, veins impressed, abaxial surface reddish-brown to light green, glabrous, resinous from short glands, veins prominent, often darker or reddish. Inflorescences from upper leaf axils, of 1-3 flowers, equaling or slightly exceeding the sub- tending leaves: peduncles terete, slender, curved, 0.4-2.0 cm long, ca 1 mm in diameter, darker green or reddish to brown, glabrous, glandular- resinous, usually verrucose; bracts obsolete or caducous, when present filiform or linear, 1-5 mm long, ca 1 mm wide, dark green or reddish, glabrous; pedicels terete or ribbed toward apex, 0.5-2.3 cm long, ca 1 mm in diameter, dark green, reddish to brown, glabrous; floral tube narrowly turbinate or obconic, 2.0-2.5 mm long, ca 2 mm wide, reddish or green, glabrous, resinous, 10- costate; calyx lobes much reduced or when present 5(or S), erect or incurved, connate at apex of ovary for less than 1 mm, each lobe filiform to narrowly triangular, 2-8 mm long, 0.5-2.5 mm wide at base, margin entire, resinous, midvein occasionally keeled, inside sulcate, green or red- dish, glabrous, glandular-resinous; corolla tube ventricose, arcuate, 1.4-2.7 cm long, 2.5-4.0 mm wide at base, broader at the middle and narrowing slightly at the oblique mouth, outside red from a yellowish base, darker at the mouth, glabrous, resinous, inside pink or yellow, glabrous, limb 54 SMITHSOXIAN COSTRIBUTIOSS TO BOTAXY 5-lobed, upper lobes rotund, erect to porrect, ca 2 6a. Gesneria cubensis var. cubensis mm long-and wide, glandular, lateral and basal lobes triangular, reflexed, ca 2 mm long and wide, glandular, margins subentire to crenate; stipitate- glandular; stamens 4, adnate to corolla base for less than 1 mm, exserted up to 2 cm, filaments curved, 2.64.3 cm long, less than 1 mm wide, reddish, glabrous, anthers rotund to oblong, 1-2 mm long, ca 1 mm wide, coherent in 2 pairs, often separating at anthesis, staminode to 6 mm long; ovary inferior, apex pilose, disc cupulate or annu- late, 5-angled, style curved, equaling the length of the stamens and exserted with them, reddish, sparsely pilose, stigma clavate to stomatomorphic or bilobed. Capsule obovoid to turbinate or globose, 3-7 mm long, 3-6 mm wide, brown to gray, smooth to verrucose, costae 5 or 10, prominent or subalate; seeds narrowly rhombic or fusiform, striated, twisted, less than 1 mm long, brown. DrscussroN.-Gesneria cubensis varies considera- bly over its range. The taxa included here have traditionally been distinguished by characters of leaf margin, minor differences in length of corolla tube, exsertion of stamens and style, amount of resin on vegetative and floral parts, stamen conna- tion, etc. These characters are probably dependent on the habitat, exposure to sun and wind, or on the stage of floral development at time of collection. In other sections of Gesneria, leaf-margin characters and leaf size and shape have been found extremely variable within one species (e.g., Gesneria acaulis and G. duchartreoides. All of the specimens in- cluded here have a similar corolla color, size, and shape suited for hummingbird pollination. From observations in the greenhouse of other species of Gesneria, the variations in lengths of stamens and style and their exsertion have been seen to vary in a single flower during anthesis. One remarkable variation is the lack of calyx Gesneria cubensis (Decaisne) Baillon, Hist. Plant. 10:60, 1888.-Morton in Leon & Alain, F1. de Cuba 43456, 1957. Pentarhaphia verrucosa Decaisne, Ann. Sci. Nat. Bot. ser. 3, 6:107, 1846.-Lindl. & Paxt. in Paxton?s F1. Gard. 3:30, pl. 250, 1852-53. [Type-collection: Mt. Liban, Cuba, J. Linden 1841 (P, holotype; BM, BR, G, K, NY, P, W, isotypes).] Pentarhaphia cubensis Decaisne, Ann. Sci. R?at. Bot. ser. 3, 6:108, 1846; F1. Serres Jard. Eur., ser. 1, 3: pl. 297, 1847 [cum var. f i corollae limbo regulari Lemaire1.-Walp., Repert. Bot. Syst. 6:736, 1847.--U?. Hook., Bot. Mag. 81: pl. 4829, 1855. Conradia verrucosa Scheidweiler in Otto & Dietrich, Allg. Gartenz. 153226, 1847 [ex char., excluding locality]. Gesneria uerrucosa (Decaisne) 0. Kuntze, Rev. Gen. 413, 1891.-Morton in Leon & Alain, F1. de Cuba 43456, 1957. Gesneria dolichostyla Urban, Symb. Ant. 7:540, 1913. [Type- collection: Las Canitas, Dominican Republic, ,%I. Fuertes 1926 (K, lectotype; A, BM, BP, E, F, G, GH, NY, P, U, W, Z, isolectotypes).] Gesneria heteroclada Urban, Repert. Spec. Nov. Regni Veg. 18:372, 1922. [Type-collection: Cape Samani, Dominican Republic, TV. Abbott 1173 (US , lectotype).] Gesneria domingensis Urban, Repert. Spec. Sov. Regni Veg. 21 :223, 1925. [Type-collection: prope Constanza, Dominican? Republic, H . tion Tuhrckheim 2023, not seen, ex char.] Gesneria scopulorum Urban PC Ekman in Urban, Ark. Bot. 224 (10):73, 1929. [Type-collection: hforne Saut d?Eau, Haiti, E. Ekman H5503 ( S , holotype, EHH, IJ, S, US, iso- Gesneria subalata Urban & Ekman in Urban, Ark. Bot. 22.4 (10):74, 1929. [Type-collection: Massif des Matheux, I?Arcahaie, Morne Delpech, Haiti, E. Ekman H9328 (S, holotype; G, IJ, K, NY, S, isotypes).] Gesneria mornincola Urban g; Ekman in Urban, Ark. Bot. 22A (l0):75, 1929. [Type-collection: Massif du Sord , Gros- Morne, Morne Chabre, Haiti, E. Eknzan H5003 ( S , holotype; EHH, IJ, S, US, isotypes).] Gesneria calcicola Alain, Mem. New York Bot. Gard. 21 (2): 144, 1971. [Type-collection: Loma del Puerto, Dominican Republic, Alain Liogier 15537 ( N Y , holotype; GH, IJ, NY, P, RDJ, US, isotypes).] Gesneria bonaoana Alain, Phytologia 253275, 1973. [Type- collection: Lorna Peguera, Bonao, Dominican Republic, Alain Liogier 17372 (XY, holotype; U S , isotype).] types).] lobes in the taxon separated in the following key. In other characters G. c. var. truncata clearly Leaves often conferted, elliptic to obovate or shows a close relationship to the typical variety. Shrubs: stems to 3 m tall. oblanceolate, 1.1-6.0 cm long, 0.5-2.3 cm wide. Key to the Varieties of Gesneria cubensis 1. Calyx lobes obvious, 2-8 mm long; Cuba, Haiti, and Dominican Republic: Provinces of Azua, Santiago, La Vega, and SamanA 6a. var. cubensis 1. Calyx lobes much reduced, less than 0.5 mm long; Dominican Republic: Province of Barahona 6b. var. truncata, new rank NUMBER 29 55 Calyx lobes obvious, extending beyond apex of floral tube, 2-8 mm long. Capsule obovoid to turbinate or globose, costae prominent. TYPE-CoLLECTIoN.-Pinal de Nimanima, St. Yago de Cuba, J. Linden 2076 (P, holotype; BM, BR, G, GOET, K, NY (Figure 32), P, TCD, W, isotypes). var. cubensis is a wide-ranging variety found in eastern Cuba and in Hispaniola growing usually on exposed limestone cliffs or in thickets, from sea level to 1300 m elevation (Figure 33). Flowering specimens have been collected during all months except April and May. I)ISTRIBUTION A D EcoLocy.-Gesneria cubensis SPECIMENS ExAMINED.-CUBA. PROVINCE OF ORIENTE: Pinal de Nimanima, St. Yago de Cuba, 1843-1844, J. Linden 2076 ([August 18441 P, holotype of Pentarhaphia cubensis Decaisne; BM, BR, G 2 sheets, GOET, K 4 sheets, NY 2 sheets, P 3 sheets, TCD, W); Bayate in decliv. mont. calcar. Picote, 14 March 1918, E . Ekman 9146 ( S ) ; Picote, 14 November 1919, E. Ekman 10110 (NY, S ) ; Sierra de Nipe, in monte calcareo Picote, 15 July 1916, E . Ekman 7384 ( S 2 sheets): farallones de La Picote, near Palmarito de Cauto, 15 September 1956, M . Lopez F . 2838 (US 2 sheets); Monte Picote, a foothill a t the southern end of Sierra de Nipe, near Palrnarito del Cauto, 400 rn, 29 January 1956, C. Morton 9714 (US); Sierra de Nipe in ?manacales? ad Rio Piloto, ca 350 m, 14 March 1915, E . Ekman 5022 ( S ) ; Florida Blanca, Alto Songo, 450 m, 10 Janu- ary 1960, Alain Liogier, J. Acuna & M . Lopez F . 7406 (US); Florida Blanca, 9 February 1957, M . Lopez F . 2850 (BH, US); mogote peak, W of Santiago, 27 February 1930, J. Roig 5054 (LS); Loma del Gato and vicinity, Cobre range of Sierra Maestra, 11 July-14 August 1925, Bro. Edmundo 7 (LS, NY); El Cuero, 400 m, 18, 19 March 1912, A?. Brit ton & J. Cowell 12771 (NY, US); Guantanamo, Monte Libanon, San Fernan- dez, ca 700 m, 24 December 1919, E. Ekman 10270 ( S ) ; Mt. Liban, 1843-1844, J. Linden 1841 (Dune 18441 P, holotype of Pentarhaphia verrucosa Decaisne; BM 2 sheets, BR, G, K 2 sheets, NY, P 2 sheets, W); Potosi, Mt. Liban, 14 March, La Luisa in Monte Toro, 9 April, and Potosi iq Monte Toro, 1 May, 1860-1864, C. Wright 3076 (GH); Loma del Esparto, Cajobabo, 31 December 1959, Alain Liogier & M . Lopez F. 7105 (US). LOCALITY UNKNOWN: ?Cuba,? no date, R. de la Sagra 816 (P); ?Cuba Orientali,? 1861, C. Wrigh t 382 ( S ) ; ?Cuba,? 1860-1864, C. Wright 3076 (EM, G 2 sheets, K , MO, P, S, W); ?Cuba,? C. Wrigh t 3076 ([1856-571 GOET, [I8611 GOET, [1865] US, no date NY). HISPANIOLA. HAITI. De- partement de 1?Artibonite: Massif du Nord, Gros-Morne, Morne Chabre, ca I125 m, 5 October 1925, E . Ekman H5003 ( S , holotype of Gesneria mornincola Urban & Ekman; EHH, IJ, S, US, isotypes); Massif des Matheux, l?Arcahaie, Morne Delpech, ca 1200 m, 14 November 1927, E . Ekman H9328 ( S , holotype of Gesneria subalata Urban; G , IJ, K, NY, S, iso- types). Departement de I?Ouest: Massif des Matheux, Mireba- lais, Morne Saut d?Eau, ca 800 m, 6 February 1926, E. Ekman FIGURE 3P.--Isotype of Pentarhaphia cubensis Decaisne and Gesneria cubensis (Decaisne) Baillon, Linden 2076 (NY). H5503 (S, holotype of Gesneria scopulorum Urban & Ekman: EHH, IJ, S, US, isotypes). DOMINICAN REPUBLIC. Province of Azua: Cordillera Central, San Juan, southern spur of Picacho del Ingenito, ca 1200 m, 8 September 1929, E. Ekman H13511 ( S ) ; Las Canitas, 1300 m, July 1912, M . Fziertes 1926 ( K , lecto- type of Gesneria dolichostyla Urban; A, BM, BP, E, F, G 3 sheets, GH, NY, P, U, W, Z); Sierra de Ocoa, San Jose de Ocoa, Bejucal, ca 1000 rn, 3 March 1929, E . Ekman HI1755 ( K , S ) . Province of Santiago: Loma del Puerto, on the road to Puerto Plata, 700-800 m, 4 June 1969, A h i n Liogier 15537 (NY, holotype (2 sheets) of Gesneria calcicola Alain; RDJ 2 sheets, US, isotypes); Loma del Puerto, 750 m, 27 March 1972, Alain Liogier 19526 (NY, US). Province of La Vega: Cordillera Central, Loma Chinguela, ca 1300 m, 12 October 1929, E. Ekman HI3738 ( S ) ; Lorna Peguera, Bonao, 8 August 1970, Alain Liogier 17372 (NY, holotype of Gesneria bonaoana Alain; US, isotype.) Province of Samana: Los Haitises, near mouth of Barracote river, Samana Bay, 150 m, 19 March 1969, Alain Liogier 14494 (NY), 14507 (GH, NY, US); Los Haitises, La Manarlita, 2 July 1930, E. Ekman HI5523 ( S ) ; Cape Samana, Samani Peninsula, sea level to 100 m, 25 March 56 SMlTHSOKIAi% COSTRIBUTIOSS T O BOTANY 1921, TV. Abbott I173 (US, lectotype o f Gesneria heteroclada Urban), 1182 (US); peninsula de Samana, Cab0 Samana, P1a)a Madama, 9 May 1930, E. Ekman H14926 (A , E H H , F, G, GH, K, NY, S, US); peninsula de Samana, Cab0 Samana, a t El Fronto, 18 June 1930, E. Ekman H15344 (C, S ) . CULTIVATED: Hort. SOC. Gard. Chiswick, 5 September 1850, hleisner Herb. sn (XY); Hort. Van Houtte, no date, no collector (BM). DIsccssIoN.-There are essentially no differences among the many described taxa, included here in Gesneria cubensis var. cubensis in Cuba and Hispaniola, which can be defended in the light of the range of variability possible among the species. Since other species of Gesneria are known across the straits, which separate the islands of the Greater Antilles, the occurrence of Gesneria cubensis in both Cuba and Hispaniola cannot be ruled out, particularly since, in Cuba, G. cubensis occurs in Oriente Province, the closest of the provinces to Hispaniola, where G. cubensis is wide- spread in scattered populations. The fruits of this subspecies are distinctive by the large costae, which are most pronounced in the populations from Alto Songo in Cuba and Massif des Matheux in Haiti. The incurved calyx lobes, although variable in width, are sulcate on the inner surface and the midvein becomes prom- inent or keeled in the capsule. Openings at the base of the corolla similar to those reported by Alain in G. calcicola have also been found in samples labeled G. bonaoana and G. dolichostyla. Since fenestrate corollas rarely occur in Gesneria, these openings may be the result of animal predation on the nectar at the base of the corolla tube, or due to a genetic atavism to polypetaly similar to that seen in arti- ficial hybrids between Gesneria species (G-1236, G. ritrina X G. pedunculosa) and in the culti- vated polypetalous clone of Sinningia aggregata. The holotype of G. heterocladn Urban was probably at Berlin and presumably destroyed. The only known duplicate of Abbott 1173 in the De- partment of Botany, Smithsonian Institution, the depository of A4bbott?s first set of collections, is here selected as lectotype. The divaricate habit of these plants from the Samana Peninsula of Hispaniola is similar to many collections of G. cubensis from Cuba, but appears to be the most divaricate of an) population on Hispaniola. The tuberculate or verrucose branches also are more like Cuban pop- ulations than others that are closer geographically. Urban described Gesneria domingensis from a specimen that lacked flowers (uon Turckheim 3023) but was reported to have been collected from the area where G. cubensis is known today. N o examples of von Turckheinz 3023 have been located, but from the description the species be- longs here. The holotype of Gesneria dolichostyla was presumably at Berlin and no longer extant. From the numerous duplicates of Fuertes 1926, the FIGURE 33.-Distribution of Gesneria cubensis var. cubensis in eastern Cuba. NUMBER 29 57 specimen at Kew, is selected as Gesneria dolichostyla. Conradia verrucosa Scheidweiler from a cultivated plant grown by lectotype for was described de Jonghe, a grower who supplied other species of Gesneria (e.g., G. libanensis, from seeds collected by Linden in Cuba) to botanical gardens. T h e locality given by Scheidweiler, ?aus dem Orgelgebirge (Bra- silien),? is probably in error, since the description fits specimens of G. cubensis (G. verrucosa Decne.), and no species of Gesneria L. are known from Brazil. Gesneria cubensis persisted in cultivation for many years following the 1840?s and was depicted in many horticultural journals. The species is not known to be in cultivation at the present time. 6b. Gesneria cubensis var. truncata (Alain) L. Skog, new rank Gesneria truncata Alain, Brittonia 20:155, 1968. Shrubs: stems to 5 m tall. Leaves not conferted, usually elliptic, 2.5-7.0 Calyx lobes much truncated at apex of floral Capsule subglobose, costae not prominent. TYPE-COLLECTION.-La Hotte, between La Cueva and Placer Bonito, Dominican Republic, R. Howard 12283 (NY, holotype (Figure 31b); US, isotype). DISTRIBUTION AND EcoLoGY.-This variety is known only from the Province of Barahona in Dominican Republic, where it grows on slopes in forests at 900-1500 m elevation (Figure 27). A col- lection of flowering material was made in August. Fruiting specimens were collected in February. SPECIMENS ExAMIKED.-HISPANIOLA. DOMINICAN REPUBLIC. Province of Barahona: La Lanza, between Monteada Nueva and Polo, 900-1000 m, Alain Liogier 14321 (GH, NY); La Hotte, between La Cueva and Placer Bonito, 5000 ft, 1 Au- gust 1950, R. Howard I2283 (NY, holotype of Gesneria trun- cata Alain); Mt. Laho, trail between La Cueva and Placer Bonito, 4500 ft, 1 August 1950, R. Howard 12283 (US, iso- cm long, 0.7-2.7 cm wide. tube, reduced, ca 0.5 mm long. type). DIscussIoN.-The sheets of the type collection Howard 12283, are labeled as coming from either La Hotte or Mt. Laho. These names are probably variations for the same locality and probably vari- ations of La Jo, a name shown on maps prepared by the Defense Mapping Agency for a hill be- tween La Cueva and Placer Bonito. 7. Gesneria glandulosa (Grisebach) Urban Gesneria glandulosa (Grisebach) Urban, Symb. Ant. 2:373, 1901.-Morton in Leon and Alain, F1. de Cuba 4:463, 1957. Pentarhaphia glandulosa Grisebach, Cat. PI. Cub. 199, 1866. Shrubs: stems woody, erect, to ca 1 m tall, ca 5 mm in diameter at 10 cm below apex, bark red, densely villous with reddish articulated glandular and eglandular trichomes, resinous above, below gray, glabrescent, and rugose, lenticels obscure; branches few, leaf scars not prominent. Leaves alternate, scattered toward branch api- ces: petioles sulcate, 0.6-1.3 cm long, 1-2 mm wide, subcoriaceous, red, densely villous, glandular-resinous; blades elliptic, oblong or ob- lanceolate, 2.8-7.3 cm long, 1.0-2.9 cm wide, base acute, margin subentire below, serrulate above, apex acute, adaxial surface plane or slightly bul- late in older leaves, green, glabrescent with a few scattered trichomes along the impressed midvein, abaxial surface green or reddish-brown, villous with trichomes more dense along the reddish prominent veins, resinous. Inflorescences in upper axils, 3- to several- flowered, length equaling the subtending leaves: peduncles terete, 1.8-2.9 cm long, ca 1 mm in diameter, red, glandular-villous, resinous; bracts 2, lanceolate, ca 1 cm long, ca 2 mm wide, red, villous; pedicels 0.9-2.4 cm long, ca 1 mm in diameter, red, villous, resinous; floral tube shortly turbinate, 2-3 mm long, 4-5 mm wide at apex, red, glandular-villous, resinous on both sides; corolla tube cylindric, curved, subventricose at middle, 2.0-2.3 cm long, 4 mm wide at base, 5 mm wide at middle, 3 mm wide at throat, outside yellow (fide Wright), pilose and glandular with short unstalked glands, inside yellow, glabrous, limb erect or porrect, 5-lobed, lobes suborbicular, 1-2 mm long, 2-3 mm wide, upper lobes dentate, lateral and basal lobes entire, sparsely glandular; stamens 4, shortly exserted, filaments curved, ca 2 cm long, glabrous, anthers not seen; ovary inferior, apex pilose, disc annular, yellow, style ca 2.2 cm long, reddish, sparsely pilose, stigma curved, clavate. 58 SMITHSONIAN COSTRIBUTIOSS T O BOTANY FIGURE 34.-Type specimens: (I, holotype of Pentarhaphia glandulosn Grisebach and Gesneria glandulosa (Grisebach) Urban, Wright 3075 (GOET); b, holotype of G. heterochroa Urban, Ekman 8741 (S). Capsule broadly turbinate, 4-6 mm long, 6-8 mm wide, reddish and pilose to gray and glabres- cent with age, persistent calyx lobes patent, costae 5, obscure; seeds rhombic, striate, twisted, ca 1 mm long, 0.5 mm wide, dark red. TYPE-COLLECTION.-EI Yunque de Baracoa, Cuba, C. I V ~ i g h t 3075 (GOET, holotype (Figure 34a); G, GH, K, KY, isotypes). DISTRIBUTION A X D EcoLocY.-Gesneyia glandti- losa is endemic to El Yunque de Baracoa in Oriente Province of Cuba in rock crevices at ca 500 m elevation (Figure 29). UnderLvood (1905) reported that IYright, in correspondence to Asa Gray on 3 June 1861, wrote that he would spend a few days on El Yunque. This species was probably collected in flower at that time. SPECI>iLSS EX.A>fIKED.-CUB.4. PROVlSCr. OF ORIENTE: El Yunque de Baracoa, C. Tr'right 3075 ([1861] GOET, holotype of Pentarhnphin g lnndulosa Grisebach; [1860-1964] G, GH, K , SY, isotypes). DIscussIoh'.-This species, known from a single collection (Wright 3075) from El Yunque de Baracoa in eastern Cuba, appears intermediate be- tween Gesneyia salicifolia and G. PurpuTascens, NUMBER 29 59 both collected from near the type locality of G. glandulosa. The erect shrubby habit, subcoriaceous leaves, and elongated sepals are similar to G. salicifolia, but the pubescence of the stem, leaves, and inflorescence, fruit shape, and the somewhat bullate condition of the leaves, among other characters, resembles G. purpurascens. Abundant fruits appear to be present on the specimens. It is unknown whether the seeds were viable. Hybridi- zation between Gesneria salicifolia and G. pur- purascens, if brought into cultivation, may deter- mine the hybrid nature of G. glandulosa. 8. Gesneria haitiensis L. Skog Gesneria haitiensis L. Skog, Baileya 18:114, 1972. Shrubs: stems woody, erect or contorted at the base to 1.3 m tall, bark brown to gray at maturity, pith reddish; young branchlets divaricate, ex- tending out from the primary branches for 3-9 cm, 1-4 mm in diameter, wider toward the apices, glabrous, resinous, lenticels elongated. Leaves alternate, crowded at the apices of the branches, resinous: petioles sulcate from a swollen base, 2-6 mm long, ca 1 mm wide, green to brown, glandular and glabrous, except for sparse rows of hairs in the sulcus; blades elliptic to obovate, 2.8-5.2 cm long, 1.5-2.7 cm wide, coriaceous, base cuneate, margin entire, apex rounded to acute, adaxial surface dark green, nitid, glabrous, except for sparse rows of hairs along the immersed mid- vein, abaxial surface lighter green, glabrous, glandular along the prominent midvein. Inflorescences from the axils of the crowded leaves, l-flowered, length equaling the subtending leaves: peduncles very short, bearing 2 lanceolate bracts, 1.5 mm long, reddish-brown, glandular; pedicels terete, 0.8-1.2 cm long, elongating in fruit, ca 1 mm in diameter; floral tube turbinate, 1.5-2 mm long, ca 2 mm wide, reddish-brown, glabrous, glandular-scaly; calyx erect, 5-lobed, the lobes connate for less than 1 mm, aestivation valvate, each lobe sulcate, lanceolate-linear, 5-9 mm long, ca 1 mm broad, apex acuminate, margin entire, exterior surface reddish-brown, glabrous, glandular-scaly, interior surface lighter red, gla- brous; corolla tube cylindric, slightly curved from the gibbous base, 0.9-1.2 cm long, 4-6 mm wide at the base, 3-4 mm wide at the middle and 2-3 mm wide at the mouth, outer surface rosy-pink, gla- brous, but slightly glandular, interior surface lighter pink, glabrous, limb patent, 1.2-1.8 cm across, 5-lobed, each lobe orbiculate, flat, ca 7 mm long, ca 8 mm broad, rosy-pink, glabrous, margin erose to scalloped, glandular; stamens 4, sub- exserted, filaments adnate to the base of the corolla for less than 1 mm, linear, curved, ca 9 mm long, ca 1 mm in diameter, white, glabrous except for a group of hairs at the bend 2 mm above the base of the filament, anthers oblong, ca 1 mm long, coherent in 2 pairs, staminode 4 mm long, lacking mature anther; ovary inferior, apex reddish, pubescent disc sinuate-annular, thickened, ca 2 mm across, glabrous, style linear, ca 7 mm long, reddish, pilose, stigma stomatomorphic, and pilose. Capsule broadly turbinate or ovoid, 4-6 mm long, 6-7 mm in diameter, gray-brown, glabrous, costae 10, prominent; seeds fusiform, twisted, ca 1 mm long, reddish-brown. TYPE-COLLECTION.-Between Leon and Fond Cochon, Haiti, L. Skog, T . T a l p e y & D . Pfister 1616 (BH, holotype; BM, BR, C, E , F, G, GH, IJ, L, M, MO, NY, P, S, US (Figure 35a), W, Z, iso- DISTRIBUTION A D EcoLocY.-Gesneria haitiensis is known only from Departement du Sud in western Haiti (Figure 27) , where it grows on limestone cliffs overhanging rivers at an elevation of ca 330 m. This species was collected in flower in July. Although in cultivation at Cornell University (G-1366), the plants have not flowered. banks of dry Riviere Voldrogue at Bras-Sec [Bois-Sec] between Lkon and Fond Cochon, Massif de la Hotte, about 15 miles SE of Jeremie, 1100 f t , 24 July 1970, L. Skog, T. Talpey Pe D. Pfister 1616 (BH, holotype: BM, BR, C, E, F, G , GH, IJ, L, M, MO, NY, P, S, US, W, Z, isotypes). D1scussroN.-The species in this section, except for Gesneria aspera and G. haitiensis, are char- acterized by a bilabiate corolla with an oblique mouth often with stamens and style far exserted, or by a corolla with an arrect limb of lobes less than 2 mm long. Gesneria aspera and G. haitiensis are similar in having a corolla with a broad patent limb of lobes more than 5 mm long and having included stamens and style. Gesneria aspera differs in being pilose on stems, leaves and inflorescences, and having leaves 1 , 5 2 5 cm long, calyx lobes 2-4 mm long and corolla 1.1-2.2 cm long. tY Pes). SPECIMENS E X A M I N E D . - H A I T I , DEPARTEMENT DU SLJD: along 60 SMITHSONIAN COSTRIBUTIOSS T O BOTANY FIGURE %.-Type specimens: a, isotype of Gesneria haitiensis L. Skog, Skog, Talpey & Pfister 1616 (US): b, lectotype of G. harrisii Urban, Harris 8670 (F). 9. Gesneria harrisii Urban Gesneria harrisii Urban, Symb. Ant. 5:497, 1908. Large shrubs or small trees: stems woody, erect, to 3 m tall, 4 cm in diameter at base, bark sub- rugose brown to gray, glabrous, very resinous, lenticels elongated; branches scattered, terete, ca 4 mm in diameter at 10 cm below apex, green to brown, glabrous, resinous, internodes 0.2-8.7 cm long, leaf scars prominent. Leaves alternate, well separated or crowded toward branch apices: petioles slender, sulcate, 7-15 mm long, ca 1 mm wide, green to brown, glabrous, glandular-resinous; blades elliptic or oblong, rarely obovate, 3.4-1 1.0 cm long, 1.3-4.4 cm wide, coriaceous, rigid, base acute, margin entire below, crenate or dentate above, apex acute, adaxial surface plane, green, glabrous, somewhat resinous, veins impressed, abaxial surface lighter green or reddish-brown, glabrous, densely glandular-resinous, veins prominent. Inflorescences of single flowers, shorter than or equaling the subtending leaves: peduncles (in- cluding pedicel) terete, alate toward apex, 2-5 cm long, ca 1 mm in diameter, green to brown, glabrous, resinous; bracts obsolete; floral tube tiirbinate or obconic, 2-3 mm long, 2-3 mm wide at apex, green to brown, glabrous, resinous, cos- tate; calyx 5-lobed, lobes connate for less than 1 mm, filiform to narrowly lanceolate, erect, be- coming incurved in fruit, 9-16 mm long, ca 1 mm wide at base, outside keeled, green, glabrous and NUMBER 29 61 resinous, inside sulcate, green, glabrous, resinous; corolla tube cylindric, wider above middle, 12-17 mm long, ca 4.5 mm wide at throat, outside sulfur yellow, glabrous, resinous, inside yellow-green, glabrous, limb bilabiate, glabrous, resinous, mar- gin subentire to denticulate, stipitate-glandular, upper lobes erect, semiorbiculate, notched, lateral lobes reflexed, rotund, 1.5-2.0 mm long, ca 1.5 mm wide, basal lobe reflexed, triangular, ca 2 mm long and wide, notched; mouth not oblique; stamens 4, staminode 5 mm long, all adnate to base of corolla for less than 1 mm, exserted to length of upper lobes of corolla, filaments curved, 12-17 mm long, ca 0.5 mm wide at base, narrower above, yellow- green, glabrous, anthers oblong, ca 1 mm long, 0.5 mm wide, coherent in 2 pairs or free; ovary inferior, apex pilose, disc short, annular, style curved, 8-16 mm long, yellow-green, darker at base, glabrous, stigma thicker than style, bilobed, papillate. Capsule turbinate to broadly turbinate, dehis- cing from the apex, 4-5 mm long, 5-6 mm wide at apex, gray or brown, glabrous, resinous, costae 5, subalate; seeds fusiform, striate, twisted, ca 1 mm long, ca 0.3 mm wide, reddish or brown. TYPE-COLLECTIoN.-G1asgow, near Troy, Ja- maica, W . Harris 8670 (F, lectotype (Figure 35b); BM, UCWI, isolectotypes). DISTRIBUTION AND EcoLocY.-Gesneria harrisii is endemic to central Jamaica (Figure 36), where it grows on shaded limestone rocks and cliffs from ca 400-850 m elevation. This species appears to be in flower all during the year. SPECIMENS ExAMINED.-JAMAICA. PARISH OF TRELAWNY: near Troy, 1500 ft, 22 November 1905, I+'. Harris 9066 (BM, F, NY, UCWI). PARISH OF MANCHESTER: cliff, Troy to Oxford, Cockpit Country, 13-18 September 1906, N . Britton 677 (F, NY, US); vicinity of Auchtembeddie, 1750 ft, 1 December 1962, G . Proctor 22974 (BM, IJ); near Auchtembeddie, ca 1250 ft, 5 November 1964, G . Proctor 25623 (IJ); near Auch- tembeddie, 1250 f t , 15 August 1970, L. Skog & G. Proctor 1636 (BH, US); N of Mandeville, near Auchtembeddie, 1750 f t , 5 November 1964, T. Talpey 19 (BH); Glasgow, near Troy, 1400 ft, 20 April 1904, W. Harris 8670 (F, lectotype of Ges- neria harrisii Urban; BM, UCWI, isolectotypes); Norway dis- trict, 1.5 mi due SW of Craig hill P.O., ca 2300 ft, 30 March 1964, G . Proctor 24755 (BM, IJ, NY, U); Cockpit Country, ca 2 mi SW of Craig head, ca 2000 ft, 20 June 1959, G . Web- ster, J. Ellis & K . Miller 8416 (BM, G, IJ, s). PARISH OF ST. ANN: Douglas Castle district, ca 2300 ft, 28 March 1965, G . Proctor 26320 (IJ). PARISH OF CLARENDON: Peckham Woods, 2500 ft, 27 September 1912, W. Harris 11184 (BM, F, NY, US), 27 December 1917, W. Harris 12782 (F, GH, K, MO, NY, UCWI, US), 7 January 1955, G. Proctor 9770 (A, IJ, NY); Peckham Woods, north of Frankfield, 2500 ft, 23 December 1955, W. Stearn 4 (A, BH, BM, K). LOCALITY UNKNOWN: Jamaica, 1844, W . Purdie sn (K, NY). DIscussIoN.-Gesneria harrisii resembles G. cu- bensis vegetatively, but differs in corolla color and shape, and in the glabrous style. The flowers are similar in shape to those of G. jamaicensis, but are not rose or pink; the longer peduncles and the often obovate leaves of G. jamaicensis also sepa- rates these two species. Urban designated two specimens as syntypes, Harris 8670 and Harris 9066. Since the types at Berlin are no longer extant, the specimen of Harris 8670 at the Field Museum of Natural History, annotated by Urban, is designated here as lectotype. 10. Gesneria heterochroa Urban Gesneria heterochroa Urban, Symb. Ant. 9270, 1924.-Mor- ton in Leon S: Alain, F1. de Cuba 4:456, 1957. Gesneria clarensis var. turqziinensis Morton, Brittonia 9:19, 1957.-Morton in Leon & Alain, F1. de Cuba 43436, 1937. [Type-collection: Pic0 Turquino, Sierra Maestra, Cuba, J. AcuEa 9712 (SV, holotype; fragment, US).] Shrubs: stems woody, erect, to 2 m tall, 3-4 mm in diameter at 10 cm below apex, bark verrucose or tuberculate above from erumpent lenticels, dark green to brown, glabrous, sparingly resinous at apex; branches numerous, internodes 4-9 mm long. Leaves alternate, scattered: petioles sulcate, 4-15 mm long, ca 1 mm wide, dark green or reddish brown, verrucose, glabrous, resinous; blades lance- olate, elliptic, or obovate, 2.0-6.8 cm long, 1.0-3.2 cm wide, subcoriaceous, base acute, margin crenu- late or serrulate, revolute, occasionally reddish, apex acute to acuminate, adaxial surface plane, smooth or occasionally rough when dry, green, glabrous, veins impressed, abaxial surface reddish- brown or green, glandular-resinous, veins reddish and prominent. Inflorescences axillary near branch apices, 1- flowered, about one-half the length of the sub- tending leaves: peduncles (including pedicel) terete or subalate toward apex, 0.3-1.2 cm long, less than 1 mm in diameter, reddish, resinous; bracts 2, at base of peduncle, each linear, 2-3 mm 62 ShlITHSOSIAS CONTRIBUTIOSS TO BOTANY FIGURE 36.-Distribution of Gesneria section Pentarkaphia in Jamaica. (G. harrisii = open circles; G. jarnaicensis = solid circles.) long; floral tube narrowly obconic, ca 4 mm long, ca 2 mm wide, reddish-brown, glabrous, resinous, costae 5 ; calyx lobes 5, erect, connate at base for less than 1 mm, each linear, 5-9 mm long, less than 1 mm wide at base, reddish and resinous both sides, outside with prominent midvein, inside sulcate; corolla tube cylindric, slightly swollen at base then narrower, slightly ampliate below mid- dle, 10-15 mm long, 2-3 mm wide at base, 4-6 mm wide above, outside yellow-green to greenish-white, glabrous, glandular-resinous, inside yellow-green, glabrous, limb slightly bilabiate, 5-lobed, lobes suborbiculate, margins crenate or erose, stipitate- glandular, upper lobes erect, 2-4 mm long, 3-5 mm wide, lateral and basal lobes reflexed, 2-3 mm long and wide; stamens 4, adnate to corolla base for ca 1 mm, exserted ca 5-8 cm beyond corolla, filaments curved 1.6-2.0 cm long, jellow-green, glandular, anthers oblong, 1.0-1.5 mm long, ca 1.0 mm wide, sparsely pilose on back, staminode ca 2.5 mm long; ovary completely inferior, apex pilose, disc annular, yellow, style curved, 1 .O-2.5 cm long, pilose, stigma capitate, papillate. Capsule narrowly turbinate, 5-6 mm long, 2.0-3.5 mm wide at apex, reddish-brown, costae 5; seeds fusiform, striated, twisted, 0.5-1 .O mm long, tawny to dark brown (Figure 19a). TYpE-CoLLECTros.-Firmeza, Sierra Maestra, Cuba, E. Ekman 8741 (S, holotype, Figure 34b). DISTRIBUTION AND EcoLocY.-Gesneria hetero- chroa is known from Oriente Province of Cuba (Figure 29) at elevations from 750 to 1830 m. I ts habitat is among rocks, on cliffs or in thickets in exposed situations. Flowering material has been collected from October to January. SPECIMEAS ExA\II~ED.-CUBA. PRO\INCE OF ORIE\TE: top of Pic0 Turquino, S. Maestra, 1-2 August 1935, J . Acunn 6787 [ster.] (NY); Pic0 Turquino, Sieria hfaestra, August 1935, J . Roig 8; G. Bucker 5.3 [ster.] (US), 59 [ster.] (US); Pic0 del Este, Sierra Maestra, 5960 ft, 1-2 August 1935, J . Acuna 9712 [ster.] (US, holotjpe fragment of Gesnerin clarensis var. turquinensis Morton); Firmeza, Sierra hlaestra, ca 750 m, 9 Noiernber 1917, E . E k m m 8741 (S, holotjpe of Gesnerza heterochroa Urban); Gran Peidra, Sierra hfaestra, ca 1200 m, Alain Liogier, J . Acuiia PC A f . Lofiez F . 7441 (US); La Gran Piedra, supra Daiquiri, ex colon. "La Floiida," ca 850 m, 28-29 June 1914, E . Ekman 1598 ( S ) ; Sierra Maestra, prope Daiquiri, 800-1000 m, 28 October 1916, E . Ekman 8090 (NY, S), DIscvssIoiv.-Gesneria heterochroa includes G. rlarensis var. turquinensis, which was described without flolvers or fruit. The vegetative characters and the range of this latter population axe closer to G. heterochroa from Oriente Province than with G. clarensis from Las Villas Province of Cuba. The slender branches bearing well-spaced leaves acute at the apex distinguish the population at Pic0 Turquino from G. uiridiflora subsp. vz'l-icli- flora at the same locality, which has stout branches and rotund or emarginate leaves. NUMBER 29 63 11. Gesneria hypoclada Urban & Ekman Gesneria hypoclada Urban 8c Ekman in Urban, Ark. Bot. 20A (5):50, 1926. Small trees: stems woody, erect, bark verrucu- lose from erumpent lenticels and yellowish when young, becoming rugose to smooth and gray-brown with age, glabrous, apices resinous; branches numerous, terete, slender, ca 2 mm in diameter at 10 cm below apex, internodes 0.1-1.1 cm long, leaf scars prominent. Leaves alternate, spiral, congested toward the branch apices: petioles sulcate, 3-6 mm long, ca 1 mm wide, verruculose, yellow from resin, glandular; blades elliptic to obovate, 2.1-4.2 cm long, 1.0-1.4 cm wide, subcoriaceous, base nar- rowly acute, margin subentire below, above crenate or serrate, apex acute, adaxial surface plane, yellow-green from resin, glandular, midvein im- pressed, abaxial surface yellow-green, glandular, veins darker, prominent. Inflorescences axillary near branch apices, 1- flowered, equaling the subtending leaves: pedun- cles curved, terete, 6-8 mm long, less than 1 mm in diameter, yellow, glandular-resinous; bracts 2, caducous, approximate, linear, 1-3 mm long, yellow, glandular-resinous, apex apiculate; pedicels terete or subalate toward apex, 4-9 mm long, less than 1 mm wide, yellow, glandular-resinous; floral tube turbinate, ca 2 mm long, 2-3 mm wide at apex, green to yellow, glandular-resinous, costae 5; calyx Nobed, lobes connate at base for less than 1 mm, erect at anthesis becoming sinuate, each narrowly triangular, 4-10 mm long, ca 1 mm wide at base, margin chartaceous, midvein prominent outside, both sides yellow-green, glandular-resinous; corolla tube cylindric, subventricose, 0.8-1.5 cm long, ca 3 mm wide at base, ca 6 mm wide above, green to brown with darker veins, glandular- resinous both sides, limb slightly bilabiate, upper lobes erect, suborbiculate, ca 3 mm long and wide, margin crenate, stipitate-glandular, lateral and basal lobes patent, orbiculate, ca 3 mm long and wide, glandular, entire; stamens 4, adnate to corolla base, exserted only to corolla limb, fila- ments 1.3-1.5 cm long, yellow-green, glabrous, anthers oblong, ca 1.5 mm long, coherent in 2 pairs at anthesis, staminode 5 mm long; ovary inferior, apex pilose, disc annular, yellow, style 1.7-1.8 cm long, yellow-green, pilose at base, stigma papillate. Capsule turbinate, ca 4 mm long and wide at apex, brown, glandular-resinous; seeds not seen. TYPE-COLLECTION.-Morne Tranchant, PCtion- ville, Haiti, E . Eknzan H1877 (S, holotype; EHH, IJ, K, NY, S, US Figure 37a, isotypes). DISTRIBUTION A D EcoLocY.-Gesneria hypoclada is known only from the mountains of Departement de I?Ouest in southeastern Haiti (Figure 27) at elevations of 1800-1825 m, where it grows on hard limestone. This species has been collected in flower only in September. SPECIMENS ExAMINED.-HISPANIOLA. HAITI. Departement de I?Ouest: Massif de la Selle, Morne Tranchant, Petionville, 1800-1825 m, 13 September 1924, E. Ekman HI877 ( S , holo- type of Gesneria hypoclada Urban & Ekman; EHH, IJ, K , NY, S, US 2 sheets, isotypes). DrscussIoN.-Gesneria hypoclada most closely re- sembles G. heterochroa from Cuba in its one- flowered inflorescences with short peduncles bear- ing greenish flowers. The Cuban species differs in being fruticose rather than arborescent, having capsules 5-6 mm long, and petioles 1.0-1.6 cm long. This Haitian species from near Petionville is probably now extinct due to its distribution near the region of dense human population and the incessant search for wood for charcoal by rural people. 12. Gesneria jamaicensis N. Britton Gesneria jamaicensis N. Britton, Bull. Torrey Bot. Club 48: 341, 1922 [1921].-Adams, F1. PI. Jamaica 680, 1972. Small straggling shrubs: stems woody, to 3.3 m tall, 3-4 mm in diameter at 10 cm from apex, bark smooth to rough, green or brown to gray, glabrous, apex verruculose from glandular tri- chomes, very resinous, lenticels obscure; branches scattered, terete, soon becoming woody, internodes 0.1-14.0 cm long, leaf scars large, prominent. Leaves alternate, spiral, scattered or congested at branch tips: 2etioles sulcate, 4-14 mm long, 1-2 mm wide, green, glabrous, glandular and resinous; blades elliptic, oblong or obovate, occasionally falcate, usually broader above the middle, 6.0-15.4 cm long, 1.9-5.7 cm wide, rigid, coriaceous, base acute, margin entire toward the base, serrate or dentate above, apex acute to acuminate, adaxial surface plane, dark green, glabrous, nitid, 64 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 37.-Type specimens: a, isotype of Gesneria hypoclada Urban & Ekman, E k m a n H1877 (US): b, holotype of G . jamaicensis Britton, Harris 12509 (XY). glandular-resinous, veins impressed, abaxial sur- face copper-colored and green at the veins or all green, glabrous, glandular-resinous, veins promi- nent and darker. Inflorescences of I(-2) flowers, length equaling or exceeding the subtending leaves: peduncles (including pedicel) terete or subquadrangular, curved, 4.5-11.8 cm long, 1-2 mm wide at base, slender at middle, broader toward the costate apex, reddish, glabrous, glandular-resinous; bracts ob- solete; floral tube obconic or turbinate, 2-8 mm long, 2-4 mm wide, dark red, glabrous, glandular- resinous; calyx lobes 5, erect, incurved, connate for ca 1 mm, each linear to lanceolate, 7-11 mm long, 1-3 mm wide at base, outside keeled, rose, glabrous, glandular-resinous, inside sulcate, green, glabrous, glandular-resinous; corolla tube sub- cylindric, broadest at base, narrowing to middle, then wider toward mouth, 8-10 mm long, 3 mm wide at base, ca 4 mm wide at mouth, outside deep rose or crimson to rose pink, glabrous and resinous, inside lighter pink, glabrous, limb 5-lobed, gla- brous, margin erose or dentate, lobes orbiculate 3-5 mm long, 3-4 mm wide, upper lobes erect to porrect, lateral and basal lobes patent to reflexed, mouth oblique; stamens 4, staminode ca 3 mm long, all adnate to base of corolla tube, not ex- serted beyond corolla limb, filaments sigmoid- curved, broad at base, filiform above, 8-10 mm long, ca 1 nim wide at base, reddish, glabrous, anthers oblong to rotund, nodding, ca 1 mm long, and wide, coherent in 2 pairs, becoming free; ovary inferior, apex pilose, disc annular, thickened, style curved, 7-8 mm long, ca 1 mm wide, reddish, sparsely pilose at base, glabrous above, stigma bilobed, papillate. NUMBER 29 65 Capsule subglobose to obconic, 4-8 mm long, 4-6 mm wide, reddish-brown to gray, glabrous, resinous, costae 5; seeds fusiform to rhombic, striated, twisted, ca 1 mm long, ca 0.25 mm wide. TYPE-CoLLEcTIoN.-Ipswich, Jamaica, w. Harris 12509 (NY, holotype, Figure 37b; BM, F, GH, K, MO, P, PH, UCWI, US, isotypes). CHROMOSOME NUMBER.-n = 14 (Lee, 1968). DISTRIBUTION A D EcoLocY.-Gesneria jamaicen- sis is known only from the parish of St. Elizabeth in western Jamaica (Figure 36), where it grows on shady limestone ledges at 250-400 m elevation. From the wild this species has been collected in flowering condition in March, June, August, and November. A cultivated plant growing in green- houses at Cornell University (G-879) produced flowers in April. Ipswich, 7 March 1917, W. Harris 12509 (NY, holotype of Gesneriu jamaicensis N . Britton; BM, F, GH, K, MO, P, PH, UCWI, US, isotypes), ca 800 ft, 10 March 1956, G . Proctor 11732 (GH), W. Stearn 442 ( A , BH, BM 4 sheets, K, S), 4 November 1964, G . Proctor 25618 (IJ), T . Talpey 16 (BH), 16 August 1970, L. Skog & G . Proctor 1640 (BH, US); Mulgrave, 1300 ft, 14 June 1916, W . Harris 12374 (BM, F , GH, K, NY, SPECIMENS ExAMINED.-JAMAICA. PARISH OF ST. ELIZABETH: UCWI, US). DIscussIoN.-Gesneria jamaicensis has been intro- duced into cultivation at Cornell University where the T. Talpey introduction of 1964 was grown under the accession numbers G-879 and G-1400. Among the species that resemble Gesneria jamaicensis, G. harrisii can be distinguished by its yellowish corolla and erect fruticose habitat; G. scabru var. sphaerocarpa, which has been collected near the range of G. jamaicensis and has a rosy- pink corolla and a straggling, suffruticose habit, can be separated by the pubescent peduncles, calyces and corollas of the multiflowered inflorescences. 13. Gesneria lanceolata Urban & Ekman Gesneria lanceolata Urban & Ekman in Urban, Ark. Bot. 22A (10):72, 1929. Shrubs: stems woody, erect to ca 2 m tall (fide Ekman); bark smooth, gray-brown, glabrous, resin- ous toward the apex; branches numerous, terete, internodes to ca 5 cm long, leaf scars prominent on young branches, becoming obscure. Leaves alternate, scattered at branch apices: petioles sulcate, 8-15 mm long, ca 1 mm wide, glabrous, resinous; blades narrowly elliptic to ob- lanceolate, 6.7-11.0 cm long 1.5-3.3 cm wide, sub- coriaceous, base acute, margin below the middle entire, above denticulate, apex acute to acuminate, adaxial surface plane, smooth, nitid, green, glabrous-glaucescent, midvein impressed, abaxial surface lighter green or light reddish-brown, veins prominent. Inflorescences in axils of upper leaves, 2- or 3- flowered, ca one-fourth to one-half the length of the subtending leaves: peduncles terete, 3-6 mm long, ca 0.5 mm wide, glabrous; bracts 2, caducous, linear, 2-3 mm long; fruiting pedicels terete, 5-7 mm long, ca 0.3 mm wide, Flowers not seen. Capsule turbinate, ca 4 mm long, ca 3.5 mm wide toward the apex, gray-brown, glabrous, the persistent calyx lobes linear, ca 1.5 mm long, ca 1 mm wide, fruit apex erumpent, pilose, costae 10; seeds fusiform, ca 0.7 mm long, brown. TYPE-COLLECTIoN.--Massif de la Selle, Croix-des- Bouquets, Haiti, E . Ekman HT861 (S, holotype; IJ, S, isotypes). DISTRIBUTION A D EcoLocY.-Gesnerin lanceolata is endemic to the Massif de la Selle in southern Haiti (Figure 27) at ca 1000 m elevation. The only collection of this species, consisting of fruiting material, was made in March. SPECIMENS ExAMINED.-HISPANIOLA. HAITI. Departement de I?Ouest: Massif de la Selle, Croix-des-Bouquets, Badeau, at Trou-a-I?Eau, ca 1000 m, 15 March 1927, E . Ekman H7861 ( S , holotype of Gesneria lanceolata Urban & Ekman (Figure 38a); IJ, S, isotypes). DIscuSsIoN.-Since Gesneria lanceolata was de- scribed without flowers, its sectional placement is not definite. On the basis of vegetative characters, Gesneria lanceolata is probably closely related to G. cubensis but is maintained separately because of the larger leaves and short inflorescences. 14. Gesneria odontophylla Urban & Ekman Gesneria odontophylla Urban & Ekman in Urban, Ark. Bot. 24A(4):34, 1931. Shrubs: stems woody, to 1 m tall, bark smooth, brown, glabrous, resinous, lenticels obscure; branches numerous above, slender, ca 1.5-5.0 mm in diameter at 10 cm below apex, internodes 0.2- 12.0 cm long, leaf scars prominent. 66 SMITHSOXIAN CONTRIBUTIONS TO BOTANY FIGURE 38.-Type specimens: a, holotype of Gesneria lanceolata Urban & Ekman, Ekman H786I (S); b, holotype of G. odontophylla Urban & Ekman, Ekman HI0194 (S). Leaves alternate, clustered at branch apices: petioles sulcate, 1.1-3.0 cm long, ca 1 mm wide, green to yellow, glandular-resinous; blades broadly elliptic to obovate, 6.1-12.0 cm long, 3.1-7.0 cm wide, membranous, base acute, margin subentire below, serrate above, apex acute to acuminate, adaxial surface plane, smooth, light green, glabrous-resinous at the reddish veins, abaxial sur- face brownish, glabrous, resinous, veins red, prominent. Inflorescences in upper leaf axils, 3-flowered: immature peduncles ca 1.5 cm long, ca 1 mm in diameter, resinous; young pedicels ca 2 mm long; flowers not seen. Capsules turbinate; seeds not seen, TYPE-CoLLEcTIoN.-Fond Cochon, near Bras-Sec, Haiti, E. Ekman Hl0194 (S, holotype (Figure 38b); EHH, IJ, K, S, US, isotypes). DISTRIBUTION AND EcoLocu.-Gesneria odonto- phyl la is known only from southwestern Haiti (Figure 27) where i t grows on rocky, shady hill- sides at ca 300 m elevation. Specimens bearing flowers are not known; the type-collection made in June bears only dry immature inflorescences. SI?ECI?.IEP;S Ex.s?.ilhED.-HISP.~SIOL..i. HAITI. Departement du Sud: Massif de la Hotte, western group, Les Roseaux, Fond-Cochon, near Bras-Sec, ca 300 rn, 28 June 1928, E . Eknian HI0194 (S, holotype of Gesneria odontophylla Urban & Ekman; EHH, IJ, K , S, US, isotypes). DrscussIoN.-Gesneria odontophylla was de- NUMBER 29 67 scribed from specimens lacking mature flowers or fruits. The type locality is apparently the same as G. haitiensis: Fond-Cochon near Bras-Sec. An at- tempt was made in 1970 to relocate G. odonto- phylla, but no material was found. This species is included here based on the habit and vegetative characters. It may be closely related to either G. clarensis from Cuba or G. cubensis from Cuba and Hispanola. Its large leaves are distinctive, similar to G. fruticosa, known also from the type locality of G. odontophylla, but the leaves of the latter are more elliptic and glabrous. 15. Gesneria parvifolia Alain Gesneria pantifolia Alain, Mem. New York Bot. Gard. 21:145, 1971. Shrubs: stems woody, erect, up to 50 cm tall, 5 mm in diameter at 10 cm below apex, bark rough, villous near apex, becoming smooth and glabres- cent below, brownish, resinous, lenticels erumpent, dark, becoming striated; branches many, short, slender, internodes up to 5.3 cm long on rapidly growing branches, or ca 1 mm long at branch apices, leaf scars prominent. Leaves usually conferted at branch apices: petioles subsulcate, 1-3 mm long, ca 1 mm wide, reddish, villous, glandular-resinous; blades elliptic, 1.5-3.3 cm long, 0.7-1.3 cm wide, subcoriaceous, base rounded, margin serrate to dentate, apex acute or rounded, adaxial surface somewhat bul- late, green, pilose at the impressed midvein, resinous, nitid, abaxial surface brownish, pilose at the prominent reddish veins, glandular-resinous. Inflorescences in upper leaf axils of 1-3 flowers, exceeding the subtending leaves: peduncles terete, 6-10 mm long; ca 1 mm in diameter, reddish, pilose; bracts linear 5-7 mm long, ca 1 mm wide, reddish, sparsely pilose, but glandular-resinous; pedicels terete, 1-7 mm long, ca 1 mm in diameter, reddish, pilose; floral tube obconic, 2-3 mm long, 2-3 mm wide at apex, dark, red, pilose; calyx lobes 5, connate for less than 1 mm, erect, each linear to narrowly triangular, 4-7 mm long, 1.0-1.5 mm wide at base, apex acuminate, both sides red and sparsely pilose, resinous, midvein prominent; corolla tube cylindrical, ca 15 mm long, ca 3 mm wide at middle, red, pilose and glandular-resinous, inside red-orange, glabrous, but resinous, limb slightly spreading, lobes 5, rotund, 1-2 mm long, margin entire, glandular; stamens 4, adnate to corolla base for less than 1 mm, filaments linear, yellow or reddish, glabrous, anthers oblong, 1.5 mm long, coherent in 2 pairs, staminode 5 mm long; ovary completely inferior, disc annular, sinuate, 1-2 mm wide, style linear, ca 1.3 cm long, reddish, pilose, stigma capitate. Mature capsule not seen, immature capsule obovoid, 3 mm long, 3 mm wide, reddish, pilose, costae 5, obscure; seeds not seen. TYPE-COLLECTION.-sierra de Baoruco, Peder- nales, Dominican Republic, Alain Liogier 13864 (NY, holotype (Figure 39a); US, isotype). DISTRIBUTION A D EcoLocY.-Gesneria parvifolia is known only from the mountains of southern Dominican Republic (Figure 27), where it grows on limestone rocks at 800-1100 m elevation. Flowering specimens have been collected in Feb- ruary and July. SPECIMENS ExAMINFD.-HISPANIOLA. DOMINICAN REPUBLIC. Province of Pedernales: Cueva de Boucan Calice, Hoyo de Pelempito, 1100 m, 3-8 July 1971, Alain Liogier 18118 (GH, NY, US); on a knoll between Boucan Calice and Hoyo de Pelempito, Sierra de Baoruco, 800 m, 13 February 1969, Alain Liogier 13861 (NY, holotype of Gesneria ~ a r u i f o l i a Alain; US, isotype). DrscussIoN.-Gesneria parvifolia is probably a close relative of G. aspera, in which the patent limb of the cylindrical corolla, the single-flowered inflorescences, and the asperate leaves are charac- teristic. 16. Gesneria pulverulenta Alain Gesneria pulverulenta Alain, Mem. New York Bot. Gard. 21(2):146, 9 June 1971. Gesneria filisepala Alain, Phytologia 22:173, 2 December 1971. [Type-collection: Hoyo de Pelempito, Dominican Republic, Alain Liogier 18132 (NY, holotype; GH, US, isotypes).] Gesneria saxatilis Alain, Phytologia 22: 173, 2 December 1971. [Type-collection: Trail between Pedernales and Aceitial, R. Howard & E. Howard 8120 (NY, holotype; BM, GH, P, S, US, isotypes).] Subshrubs to large shrubs: stems woody, erect, up to 4 m tall, 5 mm in diameter at 5 cm below apex, bark rugose where internodes close, or smooth and striated where internodes well spaced on rapidly elongating branches, reddish-brown or green, glandular-resinous, becoming gray-brown, 68 SMITHSOSIAN CONTRIBUTIONS TO BOTANY FIGURE 39.-Type specimens: a, holotype of Gesneria paruifolia Alain, Alain Liogier 13864 (NY); b, holotype of G. puluerulenta Alain, Alain Liogier 13871 (NY). with resin flaking as a yellowish or whitish pulverulence, internodes 0.2-2.3 cm long, leaf scars prominent, pith green or red; branches few or numerous, short, in upper leaf axils, somewhat flattened, lenticels elongated, erumpent, nodes slightly swollen, axillary buds densely or sparsely covered by caducpus reddish articulated trichomes, trichomes occasionally adhering to young leaves and inflorescences. Leaves alternate, when well separated on rapidly growing branches, or approximate and conferted at branch apices; petioles subsulcate, 1-5 mm long, ca 1 mm in diameter, green or reddish, glandular- resinous; blades spathulate, obovate or elliptic, 1.0-3.4 cm long, 1.0-1.7 cm wide, coriaceous, base acute to cuneate, margin entire to coarsely crenate- dentate at the obtuse or subtruncate apex, adaxial surface smooth, nitid, dark green, glabrous but glandular, veins immersed, abaxial surface light green or brownish, glabrous, but glandular, with prominent brown or red veins. Inflorescences axillary, of I(-4) flowers, often equaling the subtending leaves: peduncles terete, 1-4 mm long, ca 1 mm in diameter, glandular- resinous; bracts 2, at apex of peduncle, filiform to narrowly lanceolate, 4-9 mm long, ca 1 mm wide, green or reddish, glabrous, glandular-resinous; pedicels terete, 3-9 mm long, ca 1 mm in diameter, green or reddish; floral tube turbinate, 2-5 mm long, 2-3 mm wide, green or reddish, glabrous, nitid, glandular-resinous; calyx erect or slightly spreading, !+lobed, lobes connate for ca 1 mm, NUMBER 29 69 aestivation valvate, each lobe linear or subulate, sulcate, 7-15 mm long, 1-2 mm wide at base, apex acuminate, both sides green or reddish, glabrous, and nitid, glandular-resinous, with a prominent keel on each lobe base; corolla (Figure 16j) tubu- lar, suburceolate, 1.3-1.6 cm long, base narrow ca 3 mm across, 4-5 mm wide below the middle then narrowing to the mouth, ca 3 mm wide; outside red or red-orange, glabrous, nitid, glandular-resinous, inside yellow, glabrous, limb slightly oblique and somewhat spreading, 5-lobed, 3-4 mm wide, lobes semiorbiculate, 1-2 mm long, red, glabrous, mar- gin entire, glandular; stamens 4, adnate to base of corolla for about 1 mm, exserted beyond mouth, at anthesis for up to 3 mm, filaments linear, 1.0- 1.3 cm long ca 1 mm in diameter, yellow-green, glabrous, anthers globose, ca 1 mm long, yellow, glabrous, coherent in 2 pairs, pollen grains iso- polar, prolate, size small (21.2-23.3 pm long at the polar axis, 11.2 pm wide at the equatorial axis), amb almost circular, tricolpate, colpi ca 21 pm long, ca 1 pm wide, sexine reticulate (Figure 18c), hetero- brochate, staminode ca 3 mm long; ovary fully inferior, apex pilose, disc sinuate-annular, yellow- green, glabrous, ca 2 mm across, style linear 8-12 mm long, ca 1 mm in diameter, yellow-green or reddish, pilose, stigma capitate, somewhat bilobed, reddish, glandular. Capsule obovate-turbinate to subglobose, 5-6 mm long, 4-5 mm wide, with 5 prominent costae; seeds fusiform to linear, striate, ca 1 mm long, reddish-brown. TYPE-COLLECTION.-Between Boucan C a k e and Hoyo de Pelempito, Dominican Republic, Alain Liogier I3871 (NY, holotype, Figure 39b). DISTRIBUTION AND EcoLocu.-Gesneria pulveru- lenta is endemic to the Sierra de Baoruco of the Barahona Peninsula of southern Hispaniola (Fig- ure 27), where it has been collected from rocky limestone cliffs and deep gorges in pine woods and thickets at elevations from 800-1300 m. In the wild, plants have been collected in flower from February to August. In the greenhouse, plants bear inflorescences all during the year. SPECIMENS ExAMINED.-HISPANIOLA. DOMINICAN REPUBLIC. Province of Pedernales: trail between Pedernales and Aceitial, 4200 ft, 8-12 August 1946, R. Howard & E. Howard 8120 (NY, holotype of Gesneria saxatilis Alain; BM, GH, P, S, US, iso- types); Aceitillar Cayo, Sierra de Baoruco, 1300 m, 9 February 1969, Alain Liogier 13672 (GH, NY); Aceitillar, 28 March 1956, J . Jirnenez 3240 (BH, E, RDJ, US); Camote, Aceitillar, 31 March 1961, E. Marcano 4518 (RDJ, US); Aceitillar, 23 March 1967, E . Marcano 5261 (BH, NY, RDJ), T. Talpey 81 (BH); Aceitillar region, 26 February 1971, Alain Liogier 17913 (NY); between Boucan C a k e and Hoyo de Pelempito, E of Aceitillar, Sierra de Baoruco, 800 m, 13 February 1969, Alain Liogier 13871 (NY, holotype of Gesneria puluerulenta Alain); Hoyo de Pelempito, 800 m, 3-8 July 1971, Alain Liogier 18132 (NY, holotype of Gesneria filisepala Alain; GH, US, isotypes). CULTIVATED: Cornell University, G-1034, 3 NO- vember 1970, L . Skog 1751 (BH). DIscussIoN.-The synonyms included in this spe- cies, Gesneria filisepala and G. saxatilis, cannot be maintained as separate species. The former was distinguished by the size of the leaves, the filiform calyx lobes, and bracts, as well as the multiflowered inflorescences. Gesneria prtlverulenta differed from G. saxatilis in its pulverulence or resin, which flakes off as a whitish powder from the vegetative parts of the plant, The size difference found in nature on all three taxa are within the limits of variation displayed by representatives in cultivation. The cultivated clone was introduced by T. Talpey in 1967 and subsequently grown in the greenhouses at Cornell University under the accession number G-1034. The variation in bract and calyx lobe shape and size is quite wide. This clone also has an inflorescence of normally a single flower; but examinations of immature flowers show the presence of reduced flower buds in the axils of the linear bracts. These buds may potentially develop to give an inflorescence of 2 or 3 flowers like that found on the holotype specimen of G. filisepala. T h e duplicate specimens of Alain Liogier 18132, the type of G. filisepala, have single-flowered inflorescences with reduced buds. Specimens of Gesneria pulverulenta have been determined and distributed as G. mornincola, a synonym of Gesneria cubensis, which differs, among other characters, in having a curved corolla tube from which the stamens and style are ex- serted more than 1 cm. 17. Gesneria salicifolia (Grisebach) Urban Gesneria salicifolia (Grisebach) Urban, Syrnb. Ant. 2373, 1901. Shrubs or small trees: stems woody, erect, 0.5- 2.5 m tall, 3-6 mm in diameter at 10 cm below apex, bark somewhat scabrous, tuberculate or verruculose from erumpent lenticels, reddish 70 SSIITHSOSIAS C O S T R I B U T I O S S TO BOTANY toward apex, brown to gray below, apex resinous; branches sparse or rarely abundant, erect, appear- ing somewhat swollen below apex, internodes 0.2- 2.5 cm long, leaf scars not prominent. Leaves alternate, at the ends of the branches, but not conferted, soon falling: petioles sulcate, 3.5-12.0 mm long, ca 1 mm wide, green to brown, glandular-resinous, verruculose; blades elongate- elliptic, oblong, oblanceolate, obovate, or spathu- late, 2.5-11.2 cm long, 0.5-2.6 cm wide, sub- coriaceous, base acute or obtuse, margin entire below, serrulate or denticulate totvard apex, re- curved, apex acute, acuminate, rounded, or emar- ginate, adaxial surface plane, smooth or slightly rough when dried, dark green to yellowish, gla- brous, nitid, veins impressed, abaxial surface reddish brown, slightly resinous, veins prominent, darker. Inflorescences numerous, cymose or subcorym- bose, in the upper axils, 1- to 4 (-6)-flowered, equaling or exceeding the subtending leaves: peduncles terete or broadened toward apex, curved upward, 4-12 cm long, ca 1 mm in diameter, green or reddish-brown, glabrous, verruculose; bracts 2, caducous, linear, 2-6 mm long, less than 1 mm wide, green or reddish-brolvn, glabrous; pedicels 0.5-1.5 cm long, ca 1 mm in diameter, green or reddish-brown; floral tube obconic, 2-4 mm long, 2-3 mm in diameter at apex, green or reddish- brown, glandular-resinous; calyx 5-lobed, lobes connate at base for ca 1 mm, each lobe erect or curved, narrowly lanceolate to linear, 0.3-1.5 cm long, 0.7-2.0 mm wide, outside with midvein prominent, inside sulcate, both sides green, yellow or reddish, glandular-resinous; corolla tube cylin- dric somewhat curved, slightly ventricose at middle, 1.4-2.5 cm long, 3-5 mm wide at base, 4-8 mm wide at middle, 3-6 mm wide at throat, outside pale yellow, pale red, scarlet or yellow-orange, glandular-resinous, inside yellow, glabrous, limb 5-lobed, lobes semiorbiculate, red or yellow, gla- brous, margins stipitate-glandular, upper lobes 1-2 mm long, 1.5-2.5 mm wide, erose, lateral lobes 1.0-1.5 mm long, 1-2 mm ivide, entire, basal lobe 1-2 mm long and wide, entire; stamens 4, shortly atlnate to base of corolla tube, included or exserted to mouth of corolla tube, filaments curved, 1.1-1.7 cm long, red or yellow, anthers oblong, 1.0-2.5 mm long, 1.0-1.5 mm wide, sparsely pilose on reverse, staminode ca 2.5 mm long; ovary inferior, apex pilose, disc annular, yellow, style curved, exserted, 2.0-2.6 cm long, green or reddish, pilose, stigma capitate, papillate. Capsule obconic, turbinate or subglobose, 3-8 mm long, 2.5-5.5 mm wide at apex, brown to gray, smooth, glabrescent, glandular, costae 5 or 10, obscure or prominent; seeds fusiform, striate, twisted, ca 1 mm long, ca 0.25 mm wide, black. 17a. Gesneria salicifolia var. ferruginea ( C . Wright) L. Skog, new rank Pentarhaphia ferrugznea C. FVright in Sauialle, Anales Acad. Ci. Med. Habana 6:317, 1869.-Sauv., ?1. Cub. 93, 1873. Gesneria ferruginea (C. Wright) Urban, Symb. Ant. 2:373, 1901.-Morton in Leon & Main, F1. de Cuba 4:459, 1957. Shrubs or small trees: stems and branches slightly swollen below apex, sparsely verruculose. Leaves broadly elliptic or oblong, base obtuse, apex acute. Flowers yellowish, calyx lobes strongly angled, greenish-yellow, 5-8 mm long, corolla pale yellow. Capsule narrowly obconic, to 8 mm long, 4-5 mm wide at apex, costae 5, prominent. TYPE-CoLLECTIoN.-Cuba, c. W?igh t 3643 (GH, holotype, (Figure 40a); K?Y, US, isotypes). Key to the Varieties of Gesneria salicifolia 1 Corolla pale )ellow, callx lobes greenish-)ellor\ ; leaies broadly elliptic or oblong, bases obtuse, capsules to 8 mm long, obconic, costae prominent; Cuba: Probince of Pinar del Rio l i a . var. ferruginea, new rank 1. Corolla pale red, scarlet, or )ello\%-orange, cal)x lobes green or reddish; leaves narronly elliptic or oboiate-spathulate, rarely broadly elliptic, bases acute; capsules obconic to sub- globose, 4-6 mm long, costae usually obscure; Cuba: Probince of Oriente. 2. Leaves narrowly elliptic, oblanceolate or oblong, not spathulate, apex acute to acuminate, calyx lobes 7-15 mm long, capsules tuibinate or subglobose I7b. var salicifolia 2. Leaies oboiate to spathulate, apex rounded to emarginate; calix lobes 3-6 mm long, capsules obconic l i c var spothulata, new kariety NUMBER 29 71 DISTRIBUTION AND EcoLocY.-Gesneria S U l i C i f O l i U var. ferruginea is known only from the Province of Pinar del Rio in western Cuba (Figure 29), where i t grows in pine forests on lateritic soil near rivers. Specimens bearing flowers or flower buds were collected in February to April, August, and October. SPECIMENS ExAMINED.-CUBA. PROVINCE OF P I N A R DEL Rio: Pan de Cajalbana, 1 October 1949, J . Acuna & Alain Liogier 15687 (SV, US 2 sheets); Loma de la Cajalbana, 1 October 1949, J . Acuna & Alain Liogier sn (IJ, US); La Cajalbana, La Palma, 4 February 1956, J . Acuna, C . Morton & Alain Liogier 20118 (US); La Cajalbana, La Palma, 8 March 1951, J . Acuna & J . Roig sn (NY); Las Puercas River, west of Cajal- bana Mts., La Palma, 8 February 1952, Alain Liogier 2360 (US); pinelands, Cajalbana, La Palma, 23 February 1954, Alain Liogier 3879 (NY, U S ) ; pinelands, Cajalbana Mts., La Palma, 1 October 1949, Alain Liogier & J . Acuna 1134 (GH, US); Loma de Cajalbana in pinetis prope amnem, 10 March 1920, E. Ekman 10500 ( S ) ; Pina de Cajalbana, in savannas at the foot of the mountains, on the edge of a branch of Rio Puercos, 28 August 1923, E. Ekman 17314 (G, NY, S 2 sheets); Pan de Cajalbana, 6 April 1915, Bro. Leon 4957 (US); on top of Cajalbana, near the waterfall, 6 April 1915, Bros. Leon 8; Charles 4957 (NY); Monte La Cajalbana, 4 February 1956, C. Morton 9825 (US); Cajalbana, 1865, C . Wright sn (K); LOCALITY UNKNOWN: Cuba, C. TVright 3643 ([1865] GH, holo- type: [no date] NY 2 sheets, US, isotypes); Cuba, C. Wright sn (S). DIscuss1oN.-None of the material of the type collection (C. Wright 3643) bears any collection data, but since all later collections of this taxon, including C . Wright sn (K), have come from La Cajalbana, this is probably the area from which the type was collected. 17b. Gesneria salicifolia var. salicif olia Gesneria salicifolia (Grisebach) Urban, Symb. Ant. 2:373, 1901.-Morton in Leon & Alain, F1. de Cuba 4:459, 1957. Pentarhaphia salicifolia Grisebach, Cat. P1. Cub. 199, 1866. Gesneria gibberosa Urban, Repert. Spec. Nov. Regni Veg. 13: 477, 1915.-Urb., Symb. Ant. 9270, 1924.-Morton in Leon & Alain, F1. de Cuba 4:459, 1957. [Type-collection: Coastal cliffs near Rio Yamuri, Cuba, J . Shafer 7838 (NY, lectotype; US, isolectotype).] Small trees: stems and branches swollen below Leaves narrowly elliptic, oblong or oblanceolate, Flowers reddish, calyx lobes 7-15 mm long. Capsules turbinate or subglobose, 4-6 mm long. TYPE-COLLECTION.-La Catalina, Cuba, c. apex, verrucose. apex acute to acuminate. Wright 3074 (GOET, holotype (Figure 41a); BM, G, GH, K, MO, NY, P, isotypes). DISTRIBUTION A D Eco~oc~.-Gesneria salicifolia var. salicifolia grows in the Oriente Province of Cuba on limestone cliffs along rivers and the sea- coast at low elevations. Collections of flowering material have been made in July and August and from November to March. SPECIMENS ExAMINED.-CUBA. PROVINCE OF ORIENTE: Faral- lones, La Catalina, 19 March 1861, C . Wright 3074 (GOET, holotype; BM, G 2 sheets, GH, K, MO, NY, P, isotypes); vicin- ity Baracoa, Bahia de Taco, 2 December 1914, E. Ekman 3697 (G, K , S , US); southern Baracoa region, on the banks of Jauco River, Jauco, 17 July-4 .4ugust 1924, Bro. Leon 11990 (NY 2 sheets, US); Los Alemanes, Rio Yumuri, Baracoa, .4u- gust 1954, J . Acuna, Pino, Alonso & T?enning 19422 (SV, US 2 sheets): paredones, Rio Yumuri, Baracoa, 4 February 1952, J . Acuna & Diuz Barreto 17355 (SV, US 2 sheets); ?10s Ale- manes,? Yumuri, Baracoa, 14 January 1956, Alain Liogier & C. Morton 5082 (GH, I J ) ; ad flum. ?Yumuri,? prope Baracoa, 25 November 1914, E. Ekman 3622 (NY, S ) ; Rio Yumuri NW of Cape Maisi, 28 August 1939, Bro. Leon 17322 (US); cliffs near the mouth of Rio Yumuri, east of Baracoa, near sea level, 13 January 1956, C . Morton & Alain Liogier 9046 (BM, US 2 sheets); Rio Yamuri [sic], 6, 12 December 1910, J . Shafer 7755 (NY, US); coastal cliffs near Rio Yamuri [sic], 6, 8 De- cember 1910, J . A . Shafer 7838 (NY, lectotype of Gesneria gibberosa Urban; US, isolectotype); Ovando Mesa, Maisi, Jan- uary 1940, Bro. Leon 17579 (GH, US); Mesa de Ovando, Maid. January 1940, Prudencio Matos in Bro. Leon 17579 (US). DIscussIox.-The holotype of Gesneria gibberosa was probably at Berlin and is no longer extant. The duplicate at New York Botanical Garden is annotated ?Det. Urban,? but not in his hand. Neither of the known sheets of Shafer 7838 has flowers, and Urban described the species without flowers, but later amplified the description after obtaining Ekman collections from the type locality at Rio Yumuri. Gesneria salicifolia was based on collections by Wright from La Catalina, but no further collec- tions are known from the type locality. Material identified as G. gibberosa is known from several areas in eastern Oriente Province of Cuba and is here included within the range of G. salicifolia (Figure 29). 17c. Gesneria salicifolia var. spathulata L. Skog, new variety Frutex. Folia obovata usque ad spathulata, 2.4-5.8 cm longa, ad bases acuta, ad margines infra 72 SlLfITHSONIAN CONTRIBUTIONS T O BOTASY FIGURE 40.-Type specimens: a, holotype of Pentarhafihia ferruginea C . Wright and Gesneria salicifolia \ar . ferruginea (C. LVright) L. Skog, new rank, Wright 3643 (GH); b, holotype of G. salicifolia tar . spathula ta L. Skog, new Tariety, Acuna 13348 (SV). medium subintegra, supra medium denticulata, ad apices rotundata vel emarginata. Flores rubelli; calycis lobi 3-6 mm longi. Capsulae obconicae, 3-4 mm longae, ad apices 2.5-3.5 mm latae. TYPE-CoLLECTIoN.-Paredones del Rio Moa, Cuba, J . Acuna 13348 (SV, holotype, Figure 40b; US, isotype). DISTRIBUTION A N D EcoLocY.-Gesneria salicifolia var. spathulata is known only from near Rio Moa in Oriente Province of Cuba (Figure 29), where i t grows at low elevations. Flowering material has been collected only in November. SPECIMENS ExAMINED.-CUBA. PROVINCE OF ORIENTE: Pare- dones del Rio Moa, hloa, 15 Xovember 1945, J. Acuna 13345' (SV, holotype; US, isotype). 18. Gesneria ventricosa Swartz Gesneria uentricosa Swartz, Prodr. 89, 1788. Shrubs or small trees: stems woody to within 30 cm of apex, erect, to 3 rn tall or more, up to 2 cm in diameter, bark rugose with long cracks, reddish to gray-grown, glabrous, resinous, lenticels whitish to brown, elongated; branches from the base or in upper axils, slightly flattened, subverrucose, to 1 cm wide at 8 cm below apex, green or red, to brown, glabrous, resinous, internodes about 0.1- 3.0 cm long. Leaves alternate or approximate: petioles sul- cate, 0.5-4.5 cm long, 1-5 mm wide, green or reddish-brown, essentially glabrous, resinous; NUMBER 29 73 FIGURE 41 .-Type specimens: a, holotype of Pentarhaphia salicifolia Grisebach and Gesneria salicifolia (Grisebach) Urban, Wright 3074 (GOET); b, holotype of Pentarhaphia triflora Grise- bach and Gesneria wrightii Urban, Wright 3072 (GOET). blades ovate, elliptic to obovate, 'oblanceolate, occasionally falcate, 4.1-22.7 cm long, 1.8-7.8 cm wide, membranous to subcoriaceous, base acute to truncate or cuneate, margin denticulate to crenate, apex acute to acuminate to rounded, adaxial sur- face dark green, glabrous or with glandular tri- chomes, glossy, epidermis sometimes flaky upon drying, abaxial surface lighter green to reddish- brown, glabrous, resinous, veins prominent. Inflorescences 1 - to many-flowered, shorter than or exceeding the subtending leaves: peduncles terete, 2.0-15.9 cm long, 1-4 mm in diameter, green to reddish-brown, glabrous, resinous, with lenticels prominent, elongated, becoming obscure in age; bracts 2, linear-lanceolate, 0.6-1.5 cm long, green to reddish, glabrous, resinous; pedicels terete, jointed or not, curved, 0.5-2.3 cm long, 1-2 mm in diameter, green or reddish-brown: floral tube narrowly turbinate, 2-4 mm long, 3-4 mm wide at the apex, green or reddish, glabrous, resinous; calyx lobes 5, barely connate at base above ovary, each lobe terete and filiform, or narrowly triangular, sulcate, or flattened and keeled, 0.4-2.1 cm long, 0.5-3.0 mm wide at the base, both sides green or reddish and glabrous or glandular; corolla tubular, curved, gibbous above the base, narrowing slightly acropetally, becoming ventricose at or above the middle, 2.1-3.2 cm long, 74 3 mm wide at the base, 6-10 mm wide at the middle, narrowing to 6 mm wide at the oblique mouth, outside glabrous, resinous, inside yellow at the base, orange toward the middle, becoming red at the mouth, glabrous, limb 5-lobed, each lobe with the margin entire to erose or dentate, stipitate- glandular, upper lobes ovate to semiorbiculate, 2-6 mm long, connate for about 2 mm, erect, lateral lobes 2-5 mm long, erect to reflexed or patent, basal lobe 2-3 mm long, erect to reflexed; stamens 4, adnate for about 1 mm to the base of the corolla tube, exserted to 3 cm beyond corolla mouth, filaments linear, curved, 3.1-5.5 cm long, about 1 mm in diameter, yellow at the base, be- coming orange-red toward the apex, glabrous or sparsely pilose, anthers globose or oblong, 1-2 mm long, ca 1 mm wide, red abaxially, glandular, resinous, coherent in two pairs by their apices, staminode to 9 mm long; ovary inferior, apex tomentose with whitish erect trichomes, disc an- nular, 5-lobed, yellow, puberulent with short glandular trichomes, style linear, curved, to 5.5 cm long, ca 1 mm in diameter, yellow or green at the base, orange or red acropetally, sparsely pilose to glabrescent, stigma stomatomorphic, papillate. Capsule turbinate to nearly spherical or cyathi- form, 0.5-1.0 cm long, 4-6 mm wide, yellowish (from resin) to reddish-brown, glabrous, resinous, dehiscing into two or four valves, costae 5 or 10, prominent or becoming alate; seeds rhombic or fusiform, about 1 mm long, reddish-brown, or dark red, surface cells raised (Figure 19i). DIscussIos.-Gesneria uentricosa is here divided into two subspecies differing in characters of the inflorescences and flowers. The subspecies are geo- graphically distinct: Gesneria ventricosa subsp. cymosa is known only from the island of St. Vin- cent in the Lesser Antilles, but the typical sub- species grows on many of the other islands in the Lesser Antilles. Both subspecies may be pollinated by the same species of hummingbird, GIaucis Izir- suta, found on the same islands. SMITHSOh-IAK COSTRIBUTIOSS T O BOTASY 18a. Gesneria ventricosa subsp. cymosa (Urban) L. Skog, new rank Pentarhaphia longiflora sensu Rolfe, Bull. Misc. Inform. 1893 (81):267, 1893 [non Liridley (1827), iiec semu Cook & Collins (1903)l. Gesneria cymosa Urban, Symb. Ant. 23372, 1901. Shrubs or trees: branches red, internodes short, 1-3 mm long. Leaves alternate to approximate: petioles 0.5- 1.3 cm long, 1-2 mm in diameter, red; blades elliptic to obovate, base cuneate. Inflorescences 1- to as-flowered, exceeding the subtending leaves: peduncles 7.2-9.8 cm long, 1-2 mm in diameter, red; bracts red; pedicels appar- ently not jointed, red; floral tube red, calyx lobes sulcate or terete, red; corolla orange-red to scarlet; stamens exserted about 1 cm beyond corolla mouth, filaments sparsely pilose; style green at the base. TYPE-CoLLECTIoh-.--St. Andrews Mountain, St. Vincent, H . Eggers 6662 (F, lectotype of Gesneria cymosa Urban (Figure 42a); A, GOET, hi, Z, isolectotypes). DISTRIBUTION A D EcoLocu.-Gesneria uentricosa subsp. cymosa is known only from the island of St. Vincent (Figure 44), growing in moist forest on steep mountainsides or near rivers at 250-600 m elevation. Flowering specimens have been col- lected from January to May and November to December. SPECIMENS ExA~iirsED.-LESSER ANTILLES. ST. \?ISCENT: St. Vincent, no date, A . Anderson sn (K, 2 sheets); wayside, Mount St. Andrews, 18 November 1945, P. Beard 13-11 (F, GH, MO, NY, S, ?US): St. Vincent, no date, G. Caley sn (G); Charlotte Parish, in forest on steep mountainside up from South Rivers, 1000 ft, 27 January 1962, G. Cooley 8233 (GH, IJ, USF): St. David Parish, at the foot of the Falls of Balaine, 31 March 1962, G. Cooley 8573 (GH, IJ, USF); in montibus St. Andrews, 1200 ft, 27 December 1889, H . Eggers 6662 (F, lectotype of Gesneria cymosa Urban; A, GOET 2 sheets, hf 2 sheets, Z, isolectotypes); in sylvestris ad Calvary, 1800 ft, 28 December 1889, H . Eggers 6662 (P, US): St. \?incent, H . Eggers 6662 (F, L, 6Cl2?I); St. \?incent, no date, Dr. Greville sn (E-GL); St. \?incent, no date, Rev. 2,. Guilding 20 (BM), sn Key to the Subspecies of Gesneria ventricosa 1. Inflorescences (1 -) many-flowered, peduncles, pedicels, and calyx reddish: pedicels unjointed; filaments sparsely pilose: St. Vincent 18a. subsp. cymosa, new rank 1. Inflorescences 2- to 4-flowered, peduncles, pedicels, and calyx green; pedicels jointed, filaments glabrous; Lesser Antilles from St. Croix to St. Lucia 18b. subsp. ventricosa NUMBER 29 75 FIGURE 42.-Type specimens: a, lectotype of Gesneria cymosa Urban and Gesneria uentricosa subsp. cymosa (Urban) L. Skog, new rank, Eggers 6662 (F); b, lectotype of Gesneria uentricosa Swartz, Ryan sn (BM). (E, K); between Three Rivers and Silver Spoon, 1-7 April 1950, R. Howard I1140 (BM, GH, IJ, NY); Spring Valley above sawmill, 1500-2000 ft, 3 March 1971; R. Howard, G. Cooley & R. Weaver, 17871 (A): mountains above Chateaubel- air River, 400-750 m, 23 April 1947, C. Morton 5299 (US); valley of north fork of Cumberland river, 400-600 m, 2-3 May 1947, C. Morton 5479 (US 2 sheets); Mt. St. Andrew, 500-600 m, 7 May 1947, C. Morton 5673 (GH, US); Mount Brisbane, 300-750 m, 20-21 May 1947, C. Morton 6027 (US); upper valley of Richmond valley, 330-540 m, 26-27 May 1947, C. Morton 6257 (GH, US); St. Vincent, no date, M . Parker sn (E-GL); St. Patrick, upper Bellwood Dist., inland from Layou, 800-1500 ft, 24 February 1965, G. Proctor 25912 (IJ); St. Vincent, March 1890, H . Smith & G. Smith 55 (BM, GH): forest generally near streams, on second growth on damp hillsides, 500-2000 ft, July, H . Smith & G. Smith 55 (NY). LOCALITY UNKNOWN: ?Martinica,? 1848, Gamier sn (TCD). DIscussIoN.-Specimens of Eggers 6662 are of two, possibly three, collections. The collection cited by Urban from St. Andrews Mountain was probably in the herbarium at Berlin, now de- stroyed. From the duplicates a lectotype must be chosen. A sheet at the herbarium of the Arnold Arboretum has been annotated as lectotype of Gesneria cymosa, but no publication of this lecto- typification has been found. Better specimens of the type-collection exist and from these a specimen at the Field Museum of Natural History annotated in Urban?s hand as type material is selected here as the lectotype. The opposite leaf arrangement in Gesneria uentricosa subsp. cymosa, which Urban used to differentiate the two subspecies (as species), was not apparent from the specimens examined. 76 S!dITHSOXIAX CONTRIBUTIONS T O BOTANY Similarities may be seen to Gesneria onacaensis Rusby from northern South America, but that species often has areolate leaves and stamens ad- nate to the corolla for about 2 mm and is con- sidered to belong to the genus Rhytidophyllum. 18b. Gesneria ventricosa subsp. ventricosa Gesneria uentricosa Swartz, Prodr. 89, 1788.-Sw., F1. Ind. Occid. 2:1028, 1800.-Lunan, Hort. Jam. 1:322, 1814.- Spreng., Syst. Veg. ed. 16. 2:839, 1825.--Urban, Symb. Ant. 2:371, 1901 [non Hort. Berol. ex Hanstein (1859)l. Pentarhaphia longiflora Lindley, Bot. Reg. 13: sub pl. 1110, 1827 [nom. superfl. (?Pentaraphia?)].-Griscb., F1. Brit. It?. Ind. 460, 1862.-Hook. f., Bot. Mag. 120: pl. 7339, 1894.- Duss, Ann. Inst. Bot.-Gkol. Colon. Marseille 3:430, 1897 [non sensu Cook 8: Collins (1903)l. Conradia uentricosa (Swartz) Martius ex G. Don, Gcn. Syst. 4:650, 1838.--A. P. de Candolle, Prodr. 7:525, 1839. Conradia uentricosa (Swartz) Martius p ?angustior A . P. dc Candolle, Prodr. 7:525, 1839. [Type-collection: Guadeloupe, G. Perrottet sn (G-DC, holotype).] Pentarhaphia lindleyana Decaisne, Ann. Sci. Nat. Bot. ser. 3, 6:103, 1846.--Walp., Repert. Bot. Syst. 63735, 1847.- Hanstein, Linnaea 34:300, 1865. [Type-collection: ?In occid.,? de Ponthieu sn (G, holotype; G, isotype).] Pentarhaphia swartzii Decaisne, Ann. Sci. Nat. Bot. ser. 3, 6:103, 1846 [nom. superfl.1.-\$?alp., Repert. Bot. Syst. 6: 735, 1847. Pentarhaphia exserta (Swartz) Decaisne, Ann. Sci. Sa t . Bot. ser. 3, 6:104, 1846 [pro parte, as to Decaisne?s description, excluding synonyms of Srvartz and Martius]. Pentarhaphia montana Decaisne, Ann, Sci. S a t . Bot. ser. 3, 6:105, 1846.--TValp., Repert. Bot. Syst. 6:736, 1847.-Han- stein, Linnaea 34:298, 1865. [Type-collection: Guadeloupe, F. I?Herminier sn (P, holotype),] Pentarhaphia Porida Decaisne, Ann. Sci. Nat. Bot. ser. 3, 6:106, pl. 7, 1846.-IValp., Repert. Bot. Syst. 6:736, 1847.- Hanstein, Linnaea 34:297, 1865. [Type-collection: Guadc- loupe, Beaupertuis sn (P, lectotype),] Pentarhaphia herminieri Decaisne, Ann. Sci. Nat. Bot. ser. 3, 6:106, 1846.--TValp, Repert. Bot. Syst. 6:736, 1847.-Han- stein, Linnaca 34:297, 1865. [Type-collection: Guadeloupr. F . I?Herminier sn (P, holotype).] Pentarhaphia uentricosa (Swartz) Hanstein, Linnaea 34:299, 1865. Gesneria longipora (Lindley) Baillon, Hist. Plant. 10:59, PI. 94, 95, 1888. Gesneria angustior (A. P. de Candollc) 0. Kuntzc, Rcv. Gcn. 2:473, 1891 [?Gesnera?]. Gesneria herrninieri (Decaisne) 0. Kuntze, Rev. Gen. 2:473, 1891 [?Gesnera?] . Gesneria lindleyana (Decaisnc) 0. Kuntzc, Rev. Gcn. 2:473, 1891 [?Gesnera?] . Gesneria montana (Decaisnc) 0. Kuntzc, Rev. Gen. 2:473. 1891 [?Gesn era?]. Pentarhaphia longifbra var. lindleyana (Decaisnc) Duss, Ann. Inst. Bot.-G&ol. Colon. Marseille 3:430, 1897. Gesneria uentricosa var. 0 ovata Urban, Symb. Ant. 2:373, 1901. [Type-collection: Saba, TV. Suringar sn, L (Hort. Lugd. Bat. No. 903:310-333), lectotype]. Gesneria uentricosa var. y obouata Urban, Symb. Ant. 2:372, 1901 [nom. superfl.]. Gesneria uentricosa forma montana (Decaisne) Stehlef, Bull. SOC. Bot. France 109:31, 1962. Ge.cneria ventricosa forma obovata (Crban) Stehle, Bull. SOC. Bot. France 109:31. 1962. Gesneria uentricosa forma ouata (Urban) Stehlie, null. SOC. Rot. Francr 109:31. 1962. Shrubs or trees: branches green, internodes about 1 cm long. Leaves alternate: petioles 0.6-2.1 cm long, 3-5 min wide: blades ovate to obovate, base acute to truncate. Inflorescences 2- to 4-flowered, shorter than the subtending leaves: peduncles 2.3-15.9 cm long, 2-4 mm in diameter, green; bracts green; pedicels jointed, green; floral tube green; calyx lobes flattened to terete, sometimes keeled, green: corolla yellow-orange to red (Figure 16d); stamens exserted to 3 cm beyond corolla mouth, filaments glabrous, pollen grains isopolar, prolate, size small (19.6-21.5 pm long at the pnlar axis, 11.1 pm wide at the equatorial axis), tricolpate, colpi 15.6-16.9 pm long, mesocolpia overlapping the dolpi, apocolpia rounded to truncate, sexine sparsely reticulate, heterobrochate, lumina scat- tered, less than 0.25 pm across, muri to 1 um or more wide (Figure 17g); style yellow at the base. TuPE-CoLLECTIoN.-Paradise Mountain, hfont- serrat, J . Ryan sn (BM, lectotype, Figure 42b). CHROMO~OME NIJMBER.-~ = 14 (Lee, 1966a). subsp. ventricosa is known from most of the islands of the Lesser Antilles from St. Croix south to St. Lucia (Figures 43, 44), growing in damp ravines or forest margins at elevations from sea level to 1200 m. Flowering specimens have been collected in all months except March and October. In the greenhouse, the subspecies appears to flower irregularly. below the dam, Caledonia Gut above Ham Bay, 20 January 1961, R. Howard 15663a (A): in wet valley, Trasholrn, near Annaly, 18 January 1961, R. Howard 15663b (4). ST. BAR- T H E L E x y : no datc, Euphrasen sn (UPS). SABA: W?eg i Moun- tain, 21 .4ugust 1947, Fr. ArnoZdo 778 (U): ?Rendez-I?ous? to road of the Mountain, 600-800 m, 26 December 1965, Fr. Arnoldo 3260 (U, US); Mountain, 600 m, 14 July 1906, I. BoZdingh 1758l3 (h?Y, U); Mountain, 600 m, 16 July 1906, DISTRIBUTION A D EcoLocu.-Gesneria ventricosa SPECIMESS ExAarrvr.o.-LESSER ASTILLES. ST. CROIX: NUMBER 29 77 N ST. BARTHELEMY w ST. CROlX S A B A 6 I I 50 krn I FIGURE 43.-Distribution of Gesneria section Pentarhaphia in the Lesser Antilles (part). (G. ventricosa subsp. ventricosa = solid circles.) (Coordinates: 18?N latitude and 63?W longitude.) I . Boldingh 1783B (U); between Hellsgate and Santa Cruz, 500-600 m, 7 May 1953, A . Stoffers 3172 (C, U); Mountain, at the ?Rendevous side,? 600-680 m, 12 May 1953, A . Stoflers 3248 (IJ, U); Mountains a t the ?Rendez Vous? side, 550 m, 12 August 1953, A . Stoffers 4255 (A, U); Mountain, 6-800 m, 23 April 1885, W. Suringar sn (L); Crespeen-?Rendez-vous,? 23 April 1885, W. Suringar sn (L, lectotype of G. ventricosa var. 0 ovata Urban). ST. EUSTATIUS: top of the Quill, 400 m, 8 June 1906, 1. Boldingh 471aB [No. 2431 (U, 2 sheets); top of the Quill, 500 m, 8 June 1906, I . Boldingh 479aB [No. 2511 (U); gap of the Quill, 350400 m, 13 July 1953, A . Stoffers 3605 (U); 14 April 1885, W. Suringar sn (L); Kant, 18 May 1885, W . Suringur sn (L, 3 sheets). ST. CHRISTOPHER (St. Kitts); ex insula St. Christopheri, India occidentalis, no date, J . Forsstron sn (S); 1788, B. Euphrasen sn ( S ) ; January 1883, H. Eggers 469b (M); 1780, Fr. Masson sn (BM); forest ravine near Sandy point, 8 September-5 Oc- tober 1901, A?. Britton & j . Cowell 124 (K, NY, US): interior slopes of the Crater, 2100-2500 ft, 22 February 1959, G. Proctor 19228 (A, IJ). MONISERRAT: Centre Hills, 500 m, G November 1944, J . Beard 404 (A, K , NY, U); below actual summit of hill on trail to Chance?s pond, 14-18 June 1950, R. Howard 11912 (A, BM, U, US); area of Lang?s Soufrikre, Paradise Estate, 5-14 January 1961, R. Howard & E. Howard 15175 (A, U); summit of English Peak, 5-14 January 1961, R . Howard & E. S. Howard 15195 (A, U); ?b. in Montserrat, Gesneria flexuosa,? no date, J. von Rohr sn (C, PH); Para- dise Mountain, [ca 17801, J. Ryan sn (BM, lectotype of Ges- neria ventricosa Swartz); near Gages Soufriere, 23 January 1907, J . Shafer 187 (F, NY, US); lower slope of Farrels Mount, 25 July 1937, F. Squire sn (A). GUADELOUPE: 1839, Beazifiertuis sn (P, lectotype of Pentarhaphia florida Decaisne); Basseterre, April 1818, C . Bertero 873 (TO); ?Guadeloupe, Balb? 1821,? C . Bertero sn (G-DC); Pitons des environs des Bains-Jaunes, 1892 [1893], Pere Duss 2375 (F, GH, MO, NY, PH, US); Lavanea-Mulets, 24 April 1902, Pere Duss 4071 (F, NY, US): 1818, C. Krauss sn (G-DC, 3 sheets): ?Guadeloupe, no date,? F . I?Herminier sn (P, 2 sheets, holotypes of Pentarhaphia herminieri Decaisne and P. montana Decaisne); Montagnes de la Guadeloupe, 18 June 1824, G. Perrottet 230 (G, P); ?Montagnes de la Guadeloupe, 1825,? 18 June 1824, G . Perrottet sn (G-DC); 21 June 1824, G. Perrottet sn (G-DC, holotype of Conradia ventricosa 0 ?angustior A. P. de Can- dolle); Antilles, Guadeloupe, 1842, G. Perrottet sn (P); Saut du Constantin, 25 April 1941, A . Questel 5111 (US); Antilles, Guadeloupe, A . Questel sn (P); Guadeloupe in vajlibus um- brosis praeruptis, 1792, Herb. M . Richard sn (P); mornes basaltiques, Houelmont, 450 m, 25 February 1936, H . Stehlte 568 (US); mornes basaltiques HouElmont an Vieux Fort, 14 September 1938, H . StehEt 565 (P). DOMINICA: in silvis et ad vias, July 1881, H . Eggers 469 (BP, BR, G 3 sheets, GH, GOET, L, M, P, U, W); in silvis et ad vias, July 1881, H . Eggers 731 (K); Goodwill Valley, 23 February 1880, H . Eggers sn (US): Layou River Valley, about v2 mi above the York Valley Estate Bridge, ca 60 m, 12 April 1964, W. Ernst 1061 (US); road to Brand Bay between Fond Baron Estate and 78 S3IITHSONIAN COSTRIBUTIOSS T O BOTANY Pichelin, ca 160-450 m, 8 July 1964, TV. Ernst 1616 (Us); Riviere Douce, 28 February 1880, H . Eggers sn (GH); rain forest on the precipitous slopes of Morne Colla Anglais, Sylvania, 610-732 m, 10-23 .4ugust 1938, IV. Hodge 835 (BM, GH, NY, US 2 sheets); Sylvania, rain forest on the precipitous slopes of Morne Colla Anglais, 610-732 m, 10 February 1940, W . Hodge 1185 (GH, US); in woodlands about South Chiltern Estate, 500 m, 27-28 February 1940, t i ? . Hodge & B. Hodge 1551 (GH, KY, US); La Ronde River, 60 m, 24 November 1964, D. A?icolson 2065 (US); Roseau to Sulfur Springs in Roseau \.?alley, 27 May-2 June 1950, R. Howard 11741 (A, U, CS); Dominica, no date J . Imray 40 (K); no date, J . lmray 87 (GOET); no date, J . Imray 193 (E, K); Dominica, 1822, k?ohaut sn [Sieber Flor. Martin. Nro. 3941 (BR, GOET, R, L, M, MO, P 2 sheets, TV 4 sheets); Dominica, no date, C. Krauss 399 (E); Soufriere, 1903, F. Lloyd f68 (NY 2 sheets); water- fall, Roseau valley, 1903, F. Lloyd 806 (NY); Point Michelle, several hundred feet above the sea, 10 May 1888, G. Ramage sn (BM, K); Belfast River Valley, St. Paul, 150 ft, 2 January 1965, C. Shillingford 191 (MO, VCTVI); 2 mi N of Soufrikre, ca 75 m, in from the road to Roseau, 30 June 19i0, L. Skog 1583 (BH, US 2 sheets); St. Joseph, banks of Layou River across from Clarke Hall, 5 July 1966, I f7 . Stern & D. Tliass- hausen 2-101 (IJ, US); St. Dominique, in silvis et ad vias, July 1927, H . imn Turckheim 5381 (L); loTver slopes of Morne Plat Pays above Bellevue along trail to Grand Bay, 23 July 1964, R . Wilbur, E . Dunn, H . Hespenheide PC D. Wiseman 7871 (F, GH, MO, S Y , US). MARTISIQUE: Plateau de Larcher (entre le camp-Balata et le Camp-Colson), 1879, 1899, Pe?re Duss 329, 4543 (NY); pitons du Carbet pitons a1 marner de l?Alme, 600-900 m, 1882, Pere Duss 330 (NY); Piton du Champ flore, December 1867, L . Hahn 362 (BM, G 4 sheets, GH, P 3 sheets, U); [Martinique, 1819-18211, Kohaut s n [Sieber Flora hfixta Xo. 4071 (E-GL, G, L, LD, XY, P); east slope of L?Aileron, hit. Pelee, 950-1100 m, 8 December 1960, G. Proctor 21750 (.4): Pitons du Carbet, 1200 m, 5 September 1937, H . Stehle 2261 (US); Piton d u Carbet, 1080 m, 5 Sep- tember 1937, H . Stehle 2637 (US); entre Fonds, St. Denis et le Morne-Vert., 650 m, 27 July 1942, H . Stehle PC 121. Stehle 5173 (US); Martinique, 1823, TVrbna sn (A4\VH). ST. LLUA: Millet, 20 June 1945, P. Beard 1107 (GH, MO, S, US); Ravine Zenon (Soufriere Valley), 800 ft, 8 Xovember 1938, H. B O X 1921 (Bhl); 0.5 mi STV of Piton Troumassee, ca 2150 ft , 4 April-12 June 1958, G. Proctor 17856 (A, BM, IJ, NY, US): Great Piton, 2000 ft or more, 23 September 1888, G. Ranage sn (BM, K); rain forest along rocky streambank at foot of Mt. Gimie, ca 1500 ft, 2 .4ugust 1959, G. Tl?ebster, J . Ellis Pi K . Miller 9fII (A, IJ, S, U, US); St. Lucia, 3 August 1959, G. It?ebster, J . Ellis & K . Afiller 9f55 (A). LOCALITY UxKSO\vN: [Lesser Antilles, 1788], B. Euphrasen s n ( S ) : 1825, M . Lambut sn (G-DC); ?In occid.,? de Ponthietr sn (G, holotype of Pentarhaphia lindleynna Decaisne; G, isotype); ex India occidentali, J . Ryan stz (C, S ) : ?diffusa vanat floribus rubris et luteis e montis,? J . Ryan sn (C); J . Ryan sn [ex Herb. Schum.] (C 5 sheets); India occid.: Jamaica, 0. Suwrt: sn ( S 2 sheets); India occidentalis: Jamaica, 1i84-1786, 0. Swart: sn (S); West Indies, 1784-1786, 0. Swartz s n ( S ) . CULTIVATED: Cornell University, 2 August 1966, L . Leva 37 (BH); Cornell FIGURE 44.-Distribution of Gesneria section Pentarhaphia in the Lesser Antilles (part). (G. ventricosa subsp. ventricosa = solid circles; G. ventricosa subsp. cynosa = open circles.) NUMBER 29 79 University, 12 September 1966, M . Stone 197 (BH); Cornell University, 6 March 1972, L. Skog 1817 (BH). DIscussIoN.-With the concurrence of W. T. Stearn (in litt.), I have selected a collection by J. Ryan as the lectotype for Gesneria ventricosa. In his protolog for this species Swartz, in 1788, listed three localities from specimens he studied in the Banksian herbarium in London-now British Museum (Natural History). He later (1800) listed collectors for two of those localities: J. Ryan for Montserrat and de Ponthieu for Dominica. There is no de Ponthieu specimen from Dominica now extant in the Banksian herbarium. Thus, by elimination, J. Ryan sn is the lectotype. The third locality for Gesneria ventricosa- Jamaica-mentioned by Swartz was repeated by de Candolle ( 1 839), Grisebach (1 862), and Urban (1901), but I believe this extension of the range is erroneous. Gesneria ven tricosa occurs on the Lesser Antilles from St. Croix south to St. Vincent and has never been accurately recorded from Jamaica. The specimens at Kew labeled Gesneria ventricosa from Jamaica are undoubtedly Gesneria exserta Swartz. The Swartz specimens at Stockholm are labeled as coming from Jamaica, but the hand- writing is not that of Swartz. Possibly the specimens were obtained while Swartz worked with Joseph Banks in London, and they are perhaps duplicates of Ryan, de Ponthieu, and others collected from the Lesser Antilles, especially since no specimens of de Ponthieu are at present in the Banksian Herbarium. Pentarhaphia porida Decaisne was based on collections by Perrottet and Beaupertuis from Guadeloupe. Leeuwenberg has selected the Beau- pertuis specimen at Paris as lectotype for the name. His selection is accepted here; the specimen is annotated in Decaisne?s hand as Pentarhaphia porida. The holotype of Gesneria uentricosa var. I3 ovata Urban may have been destroyed at Berlin- Dahlem. A duplicate at Leiden of the Suringar specimen, annotated in Urban?s hand as belonging to this variety is here selected as the lectotype. The infraspecific taxa described by Urban and others appear to be ecological variants in leaf size and shape. Growing Gesneria ventricosa subsp. ventricosa collections from Dominica and St. Kitts in different regimes of light, temperature, and humidity in the greenhouse has resulted in plants similar to those described by Urban (1901), DUSS (1897), and Stehle (1962). 19. Gesneria wrightii Urban Gesneria wrightii Urban, Symb. Ant. 2:373, 1901.-Morton in Leon 8; Alain, F1. de Cuba 4:459, 1957. PentarhaPhia triflora Grisebach, Cat. P1. Cub. 199, 1866 [non Gesneria triflora Martens & Galeotti (1842), nec Gesneria triflora W. Hooker (1847)l. Small trees: stems woody, erect, to 2.5 m tall, ca 3 mm in diameter at 10 cm below apex, bark ver- rucose, green to brown above, becoming gray be- low, glabrous, apex resinous, pulverulent below, lenticels erumpent, elongated; branches occasional, scattered, internodes 0.1-2.7 cm long. Leaves alternate, usually congested at branch apices: petioles sulcate, 2-4 mm long, ca 1 mm wide, green or reddish, glabrous, verrucose, resin- ous; blades elliptic or seldom oblanceolate, 2.3- 5.3 cm long, 0.7-1.3 cm wide, coriaceous, base and apex narrowly acute, margin sharply serrate, adaxial surface light green, smooth, glabrous, veins obscure, abaxial surface light green to brown, glabrous, resinous, veins prominent. Inflorescences in axils of upper leaves, slightly longer than the subtending leaves, 2- to 4-flowered, erect: peduncles terete, curved, 1.8-5.2 cm long, ca 1 mm in diameter, red, glabrous, resinous; bracts linear, 2-5 mm long, less than 1 mm wide, resinous; pedicels terete, curved, ca 1.5 cm long, ca 1 mm wide, wider toward apex; floral tube obconic, 2-3 mm long and wide, green or red, glabrous, resin- ous, pulverulent; calyx 5-lobed, lobes erect or slightly incurved, connate for less than 1 mm at base, linear, 6-9 mm long, ca 1 mm wide at base, red, glabrous, glandular-resinous, outside keeled at base, inside sulcate; corolla tube subventricose from a narrow base, curved, 1.1-1.4 (-2.0, fide Morton) cm long, ca 2.3 mm wide at base, 3-4 mm wide at middle, narrowing slightly at throat, both sides red, glabrous, resinous, limb 5-lobed, lobes broadly rotund, short, ca 1 mm long, ca 2 mm wide, margin entire, mouth slightly oblique; sta- mens 4, adnate to base of corolla tube for less than 1 mm, slightly exserted beyond corolla tube, fila- ments curved, ca 1.4 cm long, reddish, glandular, anthers oblong, staminode not seen; ovary in- ferior, apex pilose, disc annular, style curved ca 1.4 cm long, yellow, sparsely pilose, stigma capitate. ao ShIITHSOh'IAN C O S T R I B U T I O S S T O BOTANY Capsule subglobose, ca 5 mm long and wide, red, glabrous, resinous, costae 5, prominent; seeds not seen. TYPE-COLLECTION.-San Andre, Cuba, C. Wright 3072 (GOET, holotype, Figure 41b; Bhl, G, GH, K, MO, NY, P, isotypes). DISTRIBUTION AND EcoLocu.-Gesneria 7~rightii has been found only once in Oriente Province of Cuba (Figure 29), growing on limestone cliffs. Collections with flowers were made in October. SPECI~IENS EXAMIhED.-cUBA. PROVINCE OF ORIE~TE: San Andre, C. Wright 3072 ([1861] GOET, holotlpe of Pentar- haphia trifiora Grisebach [23 October 1860-18641 GH, [1860-18641 BXI, G 2 sheets, K , MO, NY, P, isotypes). DIscussIoN.-The leaves and habit of Gesneria wrightii resemble Gesneria binghamii Morton, but differ in the smaller size of the leaves (up to 9.9 cm in G. binghamii) and corolla (up to 2.6 cm long, fide Morton), and in having inflores- cences of 2-4 flowers, rather than single flowers. Section 2. Stenochonanthe L. Skog, new section Caules erecti vel contorti, usque ad 3 m alti, ad basim vel ex foliorum superiorum axillis rami- ficantes; folia elliptica, oblanceolata vel obovata, membranacea, planiuscula, nitida; inflorescentiae 3- to co-florae, interdum ex gemmis supernumer- ariis axillaribus ortae, folia subtentes aequantes vel excedentes; flores tubulosi, corollis sensim ex basi orem versus dilatatis, flavis aurantiacis, coc- cineis, vel roseis, staminibus inclusis; capsulae turbinatae vel fere sphaericae, costis prominenti- bus vel obscuris. TYPE-SPEcIEs.-Gesneria sca bra Swartz. DIscussIoN.-Section Stenochonanthe, composed of one species including four varieties, is probably related to the other sections containing shrubs or small trees that have flowers with tubular corollas, and i t may have evolved from an ancestor of sec- tion Pentarhaphia in response to a particular pollinator. This section remains distinct, however, due to its narrowly funnel form corolla, occasional multiple axillary buds, and included stamens. Name derived from the Greek stenos (= narrow) + chone (= funnel) + anthe (= flower). 20. Gesneria scabra Swartz Gesneria scabra Swartz, Prodr. 89, 1788, Shrubs or trees: stems woody, erect or contorted, 0.5-3.0 m tall, slender, tomentose, becoming gla- brescent, resin not abundant at apex; bark reddish- gray, lenticels elongated or not evident, pith green to whitish or pink; branches rarely from super- numerary axillary buds, scattered or clustered near the apices of stems, occasional to numerous, inter- nodes 0.5-8.5 cm long, 2-3 mm in diameter, green, becoming tawny to dark brown or dark red, sub- verrucose, scabrous-pubescent to strigose or tomen- tose, trichomes often appressed, nodes slightly swollen. Leaves alternate, clustered at branch apices: petioles sulcate, 0.4-1.5 cm long, 1.0-2.5 mm wide, green to reddish or brown, scabrous-pubescent or tomentose; blades elliptic-oblong to oblanceolate or obovate, 4.3-17.2 cm long, 2.0-4.5 cm wide, membranous, rigid, plane, base subcuneate, acute or obtuse, margin serrulate-dentate or crenate, ciliate, apex acute to acuminate, adaxial surface dark green, glossy, sparsely pilose to scabrous or strigose, with appressed trichomes more abundant near the midvein, abaxial surface lighter green, sparsely pilose to scabrous or velutinous, veins prominent. Inflorescences axillary, often more than one at a node, 3- to cc-flowered, usually only one matur- ing, or in a compound dichasium and many: peduncles terete, 2.0-10.5 cm long, 1 mm in diam- eter, pilose; bracts 2, subulate to ovate, minute or 1 mm long, 1.0-2.5 mm wide, green to tawny- reddish or reddish-brown, pilose, caducous; pedi- cels terete, 0.4-2.0 cm long, 1.0-1.5 mm in diam- eter, green to reddish-brown, pilose; floral tube turbinate or nearly spherical, 2-4 mm long, 2-4 mm wide, reddish-brown, pilose, calyx lobes 5, connate to 1 mm above ovary apex, suberect to spreading, with valvate or open aestivation, each lobe flat, linear, lanceolate, lingulate, ovate, ob- ovate, or oblong, membranous, 2.5-9.0 mm long, 1-6 mm wide, accrescent, becoming chartaceous in fruit, cuneate or broad at base, pale green to white with prominent veins or reddish-brown, margin entire or serrulate, apex acute or acumi- nate, dull red or pink, scabrous-pubescent or tomentose; corolla tubular, narrowly funnelform, curved from the slightly gibbous base, tube 1.0- 1.7 cm long, 3-5 mm wide at the middle, ca 5 mm wide at the mouth, outside crimson to rosy-pink, or bright yellow to orange-yellow, pilose with NUMBER 29 81 erect, purplish articulate trichomes, inside lighter red to pink or yellow, glabrous, limb 5-lobed, expanded slightly, ca 8 mm wide, each lobe ovate, ca 2 mm long, 2-3 mm wide, pink to red, glabrous, margin erose to entire, occasionally ciliate with glandular trichomes; stamens 4, adnate to the base of the corolla tube for 1 mm, not exserted, fila- ments linear, 0.8-1.3 cm long, less than 1 mm wide, white or yellowish, glabrous, anthers oblong to sagittate, 1.0-1.5 mm long, 1.0 mm wide, white or yellowish to blackish, coherent in two pairs by their apices until dehiscence, staminode 3-4 mm long, anther lacking or infertile; ovary inferior, covered at the convex apex by dense white articu- late trichomes, disc annular, undulating, 1 mm thick, 3 mm in diameter, whitish, glabrous or tomentose, style linear, wider toward apex, ca 1.1 cm long, less than 1 mm in diameter, yellowish or orange-red, glabrous or pubescent, stigma stomato- morphic, ca 2 mm wide. Capsule turbinate to nearly spherical, 3-7 mm long, 3-7 mm wide, red, pubescent, becoming gray- brown, glabrescent, costae 5, obscure or prominent; seeds rhombic or fusiform, slightly contorted or twisted, 0.5-1.0 mm long, ca 0.7 mm wide, tawny to dark reddish-brown. DISTRIBUTION AND EcoLocu.-Gesneria scabra oc- curs in and near the Cockpit Country of western Jamaica (Figure 45). Because of the geographical proximity and the seemingly minor differences in the plants, four taxa are considered here as varie- ties, differing in corolla color, in size, color, and margin of calyx lobes, and in shape of capsule. Although the varieties are separated geograph- ically, they may be pollinated by the same species of hummingbirds, Mellkuga minima or Trochilus polytmus, whose bills are 1.3 cm and 2.0 cm long, respectively. 20a. Gesneria scabra var. fawcettii (Urban) L. Skog, new rank Gesneria fawcettii Urban, Symb. Ant. 5:500, 1908.-Adams, F1. P1. Jamaica 681, 1972. Shrubs: stems to 1 m tall. Leaves 8-13 cm long. Inflorescences few, 4- to 5-flowered: peduncles 2-6 cm long; calyx lobes lingulate to obovate, 3-6 mm long, reddish-brown, margin serrulate; corolla bright yellow; anthers yellow. Capsule turbinate, 3-4 mm long, 3-6 mm in diameter. FIGURE 45.-Distribution of Gesneria section Stenochonanthe in Jamaica. (G. scabra var. fawcettii = solid circles; G. scabra var. scabra = open circles; G. scabra var. sphaerocarpa = open squares; G. scabra var. viridicalyx = solid squares.) 82 SMITHSOSIAS CONTRIBUTIONS TO BOTAXY Key to the Varieties of Gesneria scabra 1. Corolla crimson to rosy-pink; calyx lobes entire, linear to oblong. 2. Capsule turbinate, broadest at the apex; calyx lobes ovate to oblong, 2.5-7.0 mm long 20b. var. scabra 2. Capsule nearly spherical, broadest below the apex; calyx lobes linear to lanceolate, 4-9 1. Corolla bright yellow to orange-yellow; calyx lobes serrulate, lingulate to obovate. 3. Calyx lobes reddish-brown, 3-6 mm long; inflorescences few; capsules 3-4 m m long, 3-6 3. Calyx lobes green to white, to 8 mm long; inflorescences many; capsule 3-4 mm long, 5-6 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20d. var. uiridicalyx, new variety . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . mm long . . . . . . . 20c. var. sphaerocurpa, new rank mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20a. var. fmocettii, new rank FIGURE 46.-Type specimens: a, lectotype of Gesneria fawcett i i Urban and Gesrteria scabra var. fawcettii (Urban) L. Skog, new rank, Harris 9883 (UCIVI); b, holotype of G. scnbrn var. v i i i d i - calyx L. Skog, new variety, Proctor I5756 (US). TYPE-CoLLEcTIoN.-Woodstock, Jamaica, TV. banks, at 460 m altitude, flowering in September, Harris 9853 (UCWI, lectotype, Figure 46a; BRI, F, K, NY, US, isolectotypes). var. fawcettii occurs in Jamaica, parish Of West- moreland (Figure 45), on damp, shady, limestone but probably earlier and later. SPECIJIEPI'S ExAVINED.-JAhlAICA. PARISH OF \VESTMORE- DISTRIBUT1oN EcoLocY~-Gesneria LA\D: IVoodstock, 1400 ft, 21 September 1907, W . Harris 9553 (UCIVI, lectotype of Gesneria farvcettii Urban; Bhl, F, I(, X Y , US, iso1ectotFpes). NUMBER 29 83 DISCUSSION.-TO replace the probably destroyed holotype at Berlin, the isotype specimen at Univer- sity of West Indies is selected as lectotype. 2Qb. Gesneria scabra var. scabra Gesneria scabra Swartz, Prodr. 89, 1788.-Sw., F1. Ind. Occid. 2:1020, 1800.-Lunan, Hort. Jam. 1:322, 1814.-Urb., Symb. Ant. 2:381, I9Ol.-Adams, F1. PI. Jamaica 680, 1972. Conradia scabra (Swartz) Martius, Nov. Gen. & Sp. 3:38, 1829 & 3:196, 1832.-G. Don, Gen. Syst. 4:650, 1838.-DC., Prodr. 7:526, 1839.-Griseb., F1. Brit. W. Ind. 461, 1862. Pentarhafihia scabra (Swartz) Hanstein, Linnaea 34:308, 1865. Shrubs: stems erect, to 2.5 m tall; branches erect, internodes to 2.5 cm long, green to reddish. Leaves alternate: petioles 2-9 mm long, reddish- brown; blades elliptic, oblong to obovate, 4.3-1 3.5 cm long, 2.3-4.4 cm wide, base acute to obtuse. Inflorescences 3- to %flowered: bracts minute to 5 mm long, reddish, pilose; floral tube turbinate; calyx lobes ovate to oblong, 2.5-7.0 mm long, 2-4 mm wide, base cuneate, apex acute; corolla tube 0.8-1.6 cm long, crimson to rosy, limb margin erose to denticulate; stamina1 filaments 0.8-1.3 cm long: style glabrous. Capsule turbinate, broadest at the apex, 4-7 mm long, 5-7 mm wide, costae usually 5, prominent; seeds rhombic. TYPE-CoLLECT1ox.-India occid., Jamaica, 0. Swartz sn (S, holotype, Figure 47; B, BM, C, G-DC, LD, LINN, M, S, UPS, isotypes). FIGURE 47.-Holotype of Gesneria scabra Swartz, Swart% sn (S). DISTRIBUTION AND EcoLmu.-Gesneria sca bra var. scabra grows in western Jamaica (Figure 45) in the parishes of Hanover and St. Tames. on steeD. between Cambridge and Fern, ca 1000 ft, 4 April 1966, D. I ? moist, ;Yooded, rocky limestone hilLides or thickets at elevations from ca 160 m to 550 m. This variety Powell 1669 (IJ); Burke Mountain district, 1 mi NE of Kensington, ca 1600 ft , 15 September 1957, G. Proctor 16641 (BM, IJ); Pembroke, Flagstaff, Maroon Town, 1500 ft, 11.7. appears to flower during the entire year. 1967, H . Osmaston 5004 (UCWI). LOCALITY UNKNOWN: sPECIaiENs E ~ ~ ~ ~ ~ ~ ~ . - J A M A I c A , pARIsH OF H ~ ~ ~ ~ ~ ~ : Jamaica, W . Purdie sn (K); India occid.; Jamaica, 0. Swartz sn (holotYPe, s; isotyPes, B j BM, ci G-DC, LD, LINN, M, s, ups). Retirement, 1.5 miles southeast of Kingsvale, ca 850 ft, 10 October 1950, G . Proctor 4896 (IJ, US); Retirement Moun- tain district, 1.25 mi due SE of Kingsvale, 500-800 ft, 13 January 1963, G . Proctor 23137 (BM, GH, IJ, NY, U, US); Dolphin Head, 1600 ft, 22 May 1906, W . Harris 9249 (A, BM, F, NY, UCWI, US): summit of the Dolphin Peak, January 1844, W . Purdie sn (GH, GOET, K); summit of Dolphin summit of Dolphin Head, 1789 ft, 29 October 1952, G. Proc- tor 7273 (IJ); summit of Dolphin Head, 1750-1786 ft, 11 April 1955, G. Proctor 10050 (BM, IJ, NY); N. slopes of Bubby Hill, ca 1 mi SW of Hillsbrook, 1000-1400 ft , 29 August 1965, G. Proctor 26688 (BM). PARISH OF ST. JAMES: 2Qc. Gesneria scabra var. sphaerocarpa (Urban) L. Skog, new rank Head, 17 March 1908, N . Britton & A . Hollick 2214 (F, Sy); Gesneria sphaerocarpa Symb. Ant. 5:499, F1. Jamaica 680, Shrubs or small trees: stems erect or contorted, to 3 m tall; branches often numerous, occasionally from supernumerary buds, weak and drooping or 84 SMITHSONIAN COKTRIBUTIONS TO BOTANY ascending, internodes 2.0-8.5 cm long, green, be- coming tawny to dark brown. Leaves alternate: petioles 2-9 mm long, green to reddish; blade elliptic to oblanceolate, 2.4-15.8 cm long, 1.0-3.8 cm wide, base subcuneate to acute. Inflorescences 6- to many-flowered: bracts to 8.0 mm long, 2.5 mm wide, green to tawny-reddish; floral tube nearly spherical; calyx lobes linear to lanceolate, 4-9 mm long, 1-3 mm wide, base not narrowed, apex acuminate; corolla tube 0.8-1.5 cm long, rosy-pink, limb margin entire to erose; stamina1 filaments 0.8-1.2 cm long, pollen grains isopolar, prolate, size small (18.2-18.8 pm long at the polar axis, 9.8-11.0 pm wide at the equatorial axis), colpi 14.1-15.5 pm long, overlapping to less than 1 pm wide, amb subcircular, apocolpia rounded to truncate, sexine appearing smooth, not reticulate (Figure 17f); style puberulent. Capsule nearly spherical, broadest below the middle, 3-4 mm long, 4-6 mm in diameter, costae obscure to subprominent; seeds fusiform. TYPE-CoLLEcTIoN.-Bank of Black River, La- covia, Jamaica, w. Harris 9956 (BM, lectotype; C , F, K, NY, P, UCWI, US (Figure 48), isolecto- DISTRIBUTION AND EcoLocu.-Gesneria sca bra var. sphaerocarpa is known from Jamaica, parish of St. Elizabeth (Figure 45), on limestone cliffs, from 250-450 m altitude. This variety, like the typical variety, appears to flower all during the year; in the greenhouse flower buds are present at all times. t Y pes). SPECIMEUS ExAMINED.-JAMAICA. PARISH OF ST. ELIZA- BETH: Gorge of the Jones River, south of Ginger Hill, 800- 900 ft, 25 April 1961, K . Kianer 1750 (U), G. Proctor 22238 (BM); Ipswich, 900 ft, 13 June 1916, W. Harris 12361 (BM, F, K , MO, NY, P 2 sheets, S 2 sheets, UCWI, US); Merry- wood district, between Ipswich and Mulgrave, 1200 ft, 10 March 1956, W . Stearn 446 (A, BH, BM); along road be- tween Merrywood and hlulgrave, 1000-1250 ft, 13 May 1960, G. Proctor 20845 (BM, IJ): between Merrywood and Mul- grate, 1150 ft, 16 August 1970, L. Skog & G . Proctor 1639 (BH); between Merrywood and Mulgrave, 1000 ft, 14 So- vember 1964, T. Talpey 17 (BH); bank of Black River, Lacovia, 16 September 1907, W . Harris 9956 (BM, lectotype of Gesneria sphaerocarpa Urban; C, F, K , S Y , P, UCWI, US, isolectotypes). CLLTIJATED: Cornell University, 16 hlay 1966, M . Stone 126 (BH). DIscussIoh?.-The isotype specimen of Harris 9956 at the British Museum (Natural History) has been chosen as lectotype for this variety. The holo- type was probably at Berlin and is no longer extant. 20d. Gesneria scabra var. eiridicalyx L. Skog, new variety Arbuscula ad 3 m alta. Folia ad 17.2 cm longa. Inflorescentiae flores 6-10, pedunculis 2-6 cm longis: sepala pallide viridia vel etiam albida venis prominentibus, ad 8 mm longa; antherae atratae (fide Proctor). Capsulae turbinatae, ad 3-4 mm longae et 5-6 mm latae. TYPE-CoLLECT1oN.-Between Windsor and Tyre, north of Troy, Jamaica, G. Proctor 15756 (US, holotype, Figure 46b; BM, GH, IJ, isotypes). DISTRIBUTION AND EcoLow.-Gesneria sca bra var. uiridicalyx is found in Jamaica in the Parishes FIGURE 48.-Isolectotype of Gesneria sphaerocarpa Urban and G . scabra var. sphaerocarpa (Urban) L. Skog, new rank, Har- ris 9956 (US). NUMBER 29 of St. Elizabeth and Trelawny (Figure 45) growing on moist, wooded, rocky limestone hillsides or in sinkholes at elevations from 200-550 m. Flowers are produced in March, May, August, and December. SPECIMENS ExAMINED.-JAMAICA. PARISH OF ST. ELIZA- BETH: Cooks bottom, east of Elderslie, 1600-1700 ft, 13 May 1960, G. Proctor 20865 ( B M ) . PARISH OF TRELAWNY: Cockpit country, along track between Windsor and Tyre, north of Troy, 700-1700 f t , 28 August 1956, G. Proctor 15756 (US, holotype of Gesneria scabra var. viridicalyx L. Skog; B M , GH, IJ, isotypes); Dyers Patent, WNW of Quickstep, 30 December 1949, R. Bengry sn (IJ, US); 7 miles NNW of Quickstep, 1500 ft, J. Poulter et al. in Adams I0893 (UCWI); Paynes Patent District, WNW of Quick Step, 2 August 1949, R. Bengry sn (IJ, US). Section 3. Lachnoblaste L. Skog, new section Caules erecti, usque ad 2 m alti, ex foliorum superiorum axillis ramificantes. Folia anguste obtrullata vel oblonga, interdum falcata, mem- branacea, plana, opaca. Inflorescentiae 1- ad co- florae, avam foliis subtentis breviores vel longiores; flores tubulosi ad medium ventricosi rubro- aurantiaci, staminibus exsertis. Capsulae late tur- binatae, costis prominentibus. TYPE-SPEcIEs.-Gesneria decapleura Urban. DrscussIoN.--Section Lachno blaste, consisting of a single species from Hispaniola, may have evolved from the stock ancestral to section Pentarhaphia. This section, however, differs considerably from the members of Pentarhaphia in characters of foliage and fruit. Similarities in habit, leaf shape, indu- mentum, and fruit are seen in Gesneria (Dittan- thera) fruticosa (Linnaeus) 0. Kuntze, but corolla shape easily distinguishes that species from G. decapleura. Name derived from the Greek Zachnos (= shaggy) + blastos (= bud). 21. Gesneria decapleura Urban Gesneria decapleura Urban, Symb. Ant. 7:542, 1913.-Urb., Symb. Ant. 8:647, 1921. Shrubs or trees: stems woody, erect, to 2 m tall, to 1 cm in diameter, bark rugose, white-gray to brown, glabrescent; branches smooth, gray to white or reddish, nodes slightly swollen; buds densely covered with long sordid white articulate trichomes. Leaves alternate or approximate: petioles flat- tened to subsulcate, 2-7 mm long, 1-3 mm wide, green to brown, pilose to glabrescent; blades narrowly obtrullate to oblong, occasionally falcate, 5.4-22.2 cm long, 1.3-8.6 cm wide, membranous, plane, base cuneate, margin entire near the base, becoming grossly serrate to lobulate toward the acute or acuminate apex, adaxial surface green, dull, pilose to glabrescent, veins covered with glandular or nonglandular trichomes, abaxial sur- face lighter green, pilose with articulate trichomes at least on the prominent veins. Inflorescences from the axils of often reduced leaves, 1- to m-flowered: peduncles terete, 2.6-8.0 cm long, 1 mm in diameter, reddish becoming lighter with age, pilose; bracts 2, linear-lanceolate, 3-6 mm long, less than 1 mm wide, pilose, tri- chomes with reddish to colorless articulations; pedicels terete, 0.8-1.8 cm long, 1 mm in diameter, pilose, trichomes with reddish articulations be- coming lighter with age; floral tube broadly turbinate, 2-4 mm long, 3-5 mm wide, densely pilose with reddish, articulate trichomes; calyx lobes 5, spreading, each lobe lanceolate, 5-8 mm long, 2.0-3.5 mm wide at the base, reddish and pilose with appressed trichomes outside and in- side, 3- or 5-nerved or nerves obscure; corolla tubular, curved, 1.8-2.5 cm long, 4-6 mm in diam- eter at the gibbous base, narrowing to 3-5 mm above the base, 0.8-1.1 cm in diameter at the ventricose middle, 0.7-1.0 cm in diameter at the oblique mouth, red to orange, densely pilose out- side, inside reddish and glabrescent, limb 5-lobed, each lobe semiorbiculate with margins erose to denticulate and ciliate with glandular trichomes, upper lobes 3-4 mm long, 3-4 mm wide, connate for 1-2 mm, lateral and basal lobes 2.0-3.5 mm long, 3.0-3.5 mm wide; stamens 4, adnate to the base of the corolla tube for 1 mm, exserted to about 5 mm, filaments linear, curved, 2.2-2.6 cm long, about 1 mm in diameter, yellow to red, puberulent with glandular trichomes toward the base, glabrescent above, anthers oblong, 1 .O-1.5 mm long, 1 mm wide, coherent in 2 pairs by their apices at anthesis, becoming free, staminode 3 mm long, lacking fertile anther; ovary inferior, apex densely pilose with reddish trichomes, disc annular, yellow with glandular trichomes, style linear, curved, 2.4-3.0 cm long, less than 1 mm in diam- eter, reddish, puberulent with glandular trichomes 86 SMITHSOKIAN C O S T R I B U T I O S S TO BOTANY to glabrescent above, stigma capitate, stomato- morphic. Capsule broadly turbinate, 5-6 mm long, 5-7 mm wide, reddish to gray-brown, apex slightly erumpent, dehiscing into 2-4 valves, densely pilose to glabrescent, costae 10, prominent; seeds fusi- form, twisted, ca 0.8 mm long, 0.25 mm wide, reddish-brown. TypE-CoLLEcTIoN.-Ad Las Canitas, Dominican Republic, M . Fuertes 1860 (A, lectotype, Figure DISTRIBUTION A D EcoLocu.-Gesneria decapleura is apparently endemic to Hispaniola in the central range of mountains, occurring in the northwest in the Massif d u Nord in Haiti and to the south- east in the Cordillera Central in central Domini- can Republic (Figure 50), in wet forests on lime- stone, from 150-1400 m elevation. Gesneria deca- 49) * pleura has been collected in flower during April, June, July, and August. Possible hummingbird pollinators for this species might be Anthraco- thorax dominicus and Chloiostilbon swainsonii, whose bills are 2.4 cm and 1.9 cm long, respectively. SPECIMEKS EXAMISED.--HAITI. RTEJIENT DU S O R D : Massif d u Sord, St. Louis du Nord, mountain slope at Riviere Sambiu, ca 130 m, 14 April 1925, E . E k n a n H3809 (?4, C, E H H , IJ, K , SY, S, US); Massif du Sord, St. Louis du Sord, top of Morne Charary, ca 900 m, 25 August 1925, E. Ekrnan H-liIP (S). DOMINIC.4S REPUBLIC. AZUA PROV- ISCE: Cordillera Central, Loma Kalga de Maco, headwaters of ?irr0)?O del l?alle, 1300-1400 m, 10 June 1926, E . Ekmnn H6332 (EHH, F, G, GH, IJ, S, US); ad Las Canitas, 1300 m , July 1912, M . Fuer tes 1860 ( A , lectotype of Gesneria deca- pleura Urban); ad Las Canitas, 1300 m, July 1912, Af. Fuertes IY25 (.4, S Y , P) , DIscvss1oN.-The only known duplicate of Fuertes 1860, in the herbarium of the Arnold Arboretum, has been selected as the lectotype, since the holotype at Berlin is presumed destroyed. Section 4. Myrmekianthe L. Skog, new section> Caules erecti, usque ad 2 m alti, ad basim rami- ficantes. Folia lanceolata, elliptica vel obovata, coriacea, planiuscula, nitida, marginibus recurvatis. Inflorescentiae 1- ad w-florae, folia subtentes aequantes vel excedentes; corollae tubulosae fere cylindraceae virides flavae usque rubrae, verru- cosae, staminibus non exsertis. Capsulae turbinatae, verruculosae, costis obscuris vel prominentibus. TYPE - SPECIEs.-Gesneria duchartreoides (C. Wright) Urban. D~scuss~o~. -The plants comprising section Myrmekianthe were in the past divided into five species. From the examination of many specimens over the whole geographic range of the plants, the taxa appear to be artificial and not based on strongly distinctive morphological characters nor on geographical isolation. Since flowers are sel- dom present on preserved material (and living plants are not available), attempts were made to distinguish the taxa on the basis of bracts, sepals, capsules, and vegetative organs; none of the char- acters were dependable either singly or in com- bination. Variation appears to occur over the entire distributional range; consequently, the taxa are combined here as a single variable species. Relationships of section Myrmekianthe lie, NUMBER 29 FIGURE 50.-Distribution of Gesneria section Lachnoblaste and section Dittanthera in Hispaniola. (G. (Lachnoblaste) decapleura = open circles; G . (Dittanthera) fruticosa = solid circles.) perhaps, with section Pentarhaphia on one hand and section Duchartrea on the other. The floral characters resemble superficially those found in some members of section Pentarhaphia, but stem, foliage, and fruit characters tend toward those that appear in Gesneria (Duchartrea) viridiflora subsp. viridipora, also endemic to Cuba. Plants in section Myrmekianthe are probably hummingbird pol- linated; Duchartrea may have evolved from an ancestor of this section in response to bat pollination. (= wart) + anthe (= flower). Name derived from the Greek myrmekia 22. Gesneria duchartreoides (C . Wright) Urban Gesneria duchartreoides (C. Wright) Uyban, Symb. Ant. 2:375, 1901.-Morton in Leon & Alain, F1. de Cuba 4:457, 1957. Pentarhaphia duchartreoides C. Wright in Sauvalle, Anales Acad. Ci. Med. Habana 6:316, 1869.-Sauv., F1. Cub. 92, 1873 [?du.chartraeoides?]. Gesneria nipensis N. Britton & P. Wilson, Mem. Torrey Bot. Club 16:109, 1920.-Morton in Leon & Alain, F1. de Cuba 43457, 1957. [Type-collection: Sierra Nipe, near Wood- fred, Cuba, ]. Shafer 3201 (NY, holotype; F, US, isotypes).] Gesneria norlindii Urban, Symb. Ant. 9:268, 1924.-Morton in Leon & Alain, F1. de Cuba 43458, 1957. [Type-collection: Inter Taco et Nibujon, Cuba, E . Ekman 3730 ( S , holotype).] Gesneria bracteosa Urban, Symb. Ant. 9:269, 1924.-MOTtOn in Leon & Alain, F1. de Cuba 4:458, 1957. [Type-collection: Baracoa, Lomas de Cuaba, Cuba, E . Ekman 4296 ( S , holo- type; K, S, isotypes).] Gesneria pachyclada Urban, Symb. Ant. 9:270, 1924.-Urb., Repert. Spec. Nov. Regni Veg. 21:69, 1925.-Morton i n Leon & Alain, F1. de Cuba 4:458, 1957. [Type-collection: Sierra de Cristal, Cuba. E . Ekman 6822 ( S , holotype).] Shrubs or small trees: stems woody, erect, to 2 m tall, 5-7 mm in diameter immediately below the apex; bark yellow, gray, or reddish-brown, mi- nutely puberulent to glabrous, smooth when young, becoming verrucose with age, lenticels elongated, sometimes erumpent, apex resinous; pith reddish; branches few, from the base, inter- nodes to 2 cm long. Leaves alternate: petioles sulcate to nearly terete, 0.3-2.5 cm long, 1-3 mm wide, reddish-brown, smooth to verrucose, minutely puberulent to glabrescent, resinous; blades narrowly elliptic to broadly obovate, 3.2-17.7 cm long, 0.8-5.9 cm wide, coriaceous, base cuneate or acute, margin subentire to grossly serrate, apex retuse to acumi- nate, adaxial surface green, smooth to scabrous, finely puberulent to glabrescent, glossy, abaxial surface green to reddish-brown, glabrous, mid- vein prominent, verrucose, resinous. Inflorescences 1 - to co-flowered: peduncles terete, 2.8-22.1 cm long, 1-5 mm in diameter, green to reddish-brown, puberulent to glabrescent, verru- cose; bracts 2 (-3), linear to broadly lanceolate, 0.3-4.2 cm long, 1-8 mm wide, green to reddish- brown, puberulent, apex sometimes caudate, ver- rucose, resinous; pedicel terete, 0.3-1.2 cm long, elongating in fruit, 1-2 mm in diameter, green to SMITHSONIAS C O S T R I B U T I O N S T O BOTAKY reddish-brown, puberulent, resinous; floral tube narrowly turbinate, 3-6 mm long, 3-6 mm wide, green to yellow or red, puberulent, verrucose, resinous; calyx lobes erect, 5, each triangular, linear to lanceolate, usually sulcate at the base, narrowing toward the terete apex, 0.3-2.5 cm long, 1-4 mm wide, green to reddish-brown, puberulent to glabrous, verrucose outside, inside glandular- resinous; corolla tube nearly cylindrical, 1 .0-1.7 cm long, 3-8 mm wide at the base, slightly con- tracted, then subventricose above the base, the mouth to 7 mm wide, green to yellow or red, pilose, puberulent to glabrescent, resinous outside, inside verrucose, resinous, limb 5-lobed, each lobe semi- orbiculate, 2-3 mm long, sometimes red-spotted, verrucose, sometimes pilose, margin entire, erose to fimbriate; stamens 4, adnate to the base of the corolla tube, included, filaments linear, reddish, 1.2-1.4 cm long, 1 mm in diameter, pilose to glabrescent near the base, glabrescent toward the apex, anthers oblong, 1-2 mm long, 1.5 mm wide, white, coherent in 2 pairs by their apices; ovary inferior, apex puberulent, disc annular, yellow, pubescent with glandular trichomes, style 1.3-1.6 cm long, reddish pilose to glabrescent, stigma bi- lobed, 1.5 mm wide. Capsule turbinate to nearly spherical, dehiscing into 2-4 valves, 0.4-1.1 cm long, 0.4-1.1 cm wide, red or yellow to gray-brown, verrucose, costae obscure or prominent; seeds linear, twisted, ca 1 mm long, 0.25 mm wide, reddish to brown (Figure 19n). TYPE-CoLLECTION.-cuchillas de Baracoa, Cuba, C . W r i g h t sn (GH, holotype; GOET, isotype, Fig- ure 51). DISTRIBUTION AND EcoLocY.-Gesneria duchartre- oides is known only from Oriente Province of Cuba (Figure 52), where it grows on serpentine pine barrens, stream banks, or dry hillsides from 350-1300 m elevation. This species has been col- lected in every month, but seldom with flowers. Possible hummingbird pollinators might be Calypte helenae and Chlorostilbon ricordii, whose bills are approximately the same length as the corolla tube. SPECI?.\IENS E X A M I N E D . - c U B A 4 . ORIENTE PRO1 I s m : in ?charrascales? ad Rio Piloto, 21 July 1914, E . Ekrnan 2115 ( S ) ; Sierra de Nipe in ?charrascales? ad Rio Piloto, 20 April 1919, E. Ekman 9528 ( S Y , S); Sierra de S ipe (Salto del Sojo), July 1940, Bro. Leon 17971 (US); Sojo cascade, Sierra de Sipe, Mayari, July 1940, Bro. Leon 8s Liogier 17971 (GH); Casimba, Sierra de Nipe, 2 i July 1940, Bro. Leon 19236 (US); La Casimba, Sierra de Sipe, Mayari, 17 July 1940, Bro. Leon & Alain Liogier 19236 (GH); banks of Rio Guayabo, Sierra de S ipe , Mayari, 29 July 1941, Bros. Leon & Clemente, & R. Hou?ard 20fOR (GH); banks of Rio Guayabo, Sierra de S ipe , 29 July 1911, Bro. Leon 20408 (US); Sierra de Nipe in cacum. mont. Lorna Mensura loc. rup., ca 1000 m, 16 May 1915, E. Ekmnn 5738 (G, S); Sierra de S ipe in pinetis ad pedem mont. Loma Mensura, ca 7 2 3 m, 18 February 1918, E. Ekman 9061 ( S ) ; Cayo de Monte, S of Loma Mensura, Sierra de Sipe, 7 April 1941, Bro. Leon 19973 (US); Lorna de la Bandera, Sierra dc h?ipe, March 1944, Bros. Af. I?ictorin & Clemente in Leon 22093 (LS); bank of creek near Wood- fred village, Sierra de Nipe, June-August 1941, R. Howard 6123 ( B M , GH, S Y , P, S, US); road to ?Mina U?oodfred,? FIGURE 51 .-Isotype of Pentarhaphia duchartreoides C . LVright and Gesneria duchartreoides (C. U?right) Urban, TVright sti (GOET). NUMBER 29 89 FIGURE 52.-Distribution of Gesneria section Myrmekianthe in eastern Cuba. (G. duchartreoides = solid circles.) Sierra de Nipe, 26 July 1940, Bro. Leon 19142 (US); camino de la Mina Woodfred, Sierra de Nipe, 6 April 1941, Bro. Leon 19830 (US); along Rio Medio near Woodfred, Sierra Nipe, 500 m, 18 October 1941, C. Morton & J . Acuna 3276 (BM, F , G , GH, MO, NY, S, US 2 sheets): Sierra Nipe, near Woodfred, 450-550 m, 17 December 1909, J . Shajer 3201 (NY, holotype o f Gesneria nipensis N. Britton & P. Wilson; F, US, isotypes); ?charrascales? of Loma del Winch, Sierra de Nipe, 26 February 1956, M . Lopez F. 2573 (US); El Prado, Nicaro, Mayari, 26, 27 May 1955, J. Acuna & F. Zayas 19786 (SV, US 2 sheets); charrascales de 10s pinares de Micara, 2-7 April 1956. J. Acuna, Alain Liogier & M . Lopez F. sn (US 2 sheets); charrascales, Sierra de Micara, ?Corea? lumber camp, 27 December 1955, Alain Liogier 8: M . Lopez F. 4640 (GH, IJ, US 2 sheets); Sierra de Micara, in pinelands, ca 750 m, 13 December 1922, E. Ekman 15918 (G, LS, NY 2 sheets, S); Sierra de Micara, in pinelands, 700-850 m, 17 December 1922, E. Ekman 16020 (S); charrascales de la subida a1 Sierra Cristal, 2-7 April 1956, J. Acuna, Alain Liogier & M . Lopez F. sn (US 2 sheets); Arroyo Cristal, Sierra de Cristal, 29 December 1995, Alain Liogier & M. Lopez F . 4802 (US): Sierra de Cristal prope cacum. mont. in ?charrascales- tibisiales,? 1200-1300 m, 8 March 1916, E. Ekman 6822 (S, holotype of Gesneria pachyclada Urban); Pic0 del Cristal, Sierra del Cristal, 1900-3750 ft, 25 August 1959, M . Lopez F . 84 (US); entre Las Milis (?) y la Corea, camino de Ulayan abajo a la Sierra del Cristal, 27-25 August 1959, M. Lopez F . 273 (US); Guantanamo, Monte Libanon, San Fernandez in pinetis, ca 700 m, 24 December 1919, E. Ekman I0257 (S); Camp La Gloria, south of Sierra Moa, 24-30 December 1910, J. Shafer 8135 (SY) ; Camp La Gloria, south of Sierra Moa, 24-30 December 1910, J. Shafer 8271 (NY); Pres l?arroyo Jocotea, sud de Moa, 22 July 1941, Bros. Leon & Clemente 20153 (LS): Moa, April 1943, Bro. M . Victorin & Alain Lio- gier 21440 (LS); minas de Iberia ad Taco Bay, ca 800 m, 7-8 December 1914, E . Ekman 3813 ( S ) ; Camp La Gloria, Monte La Brena, Moa, 5 November 1945, J. Acuna 13347 (SV, US 3 sheets); Moa, Rio Yagrumajes, 14 April 1945, J. Acuna 12720 (SV, US 2 sheets); banks of Cayoguan River, Moa region, 13-14 July 1949, Alain Liogier & Bro. Clemente 899 ( U S ) ; Mina Aromita [?I, Cayoguan, Punta Gorda, 24 July 1944, Bro. Clemente, Alain Liogier & Bro. Chrysogone 4066 (US); valley of Cayoguan river, Moa, 30 March 1942, Bros. Leon, M . Victorin & Ckmente 20931 (US); thickets, El COCO, Moa region, August 1945, Bros. Leon, Clemente & Alain Liogier 22639 (US); Pinal de Pena Prieta, 22 July 1953, Alain Liogier 3181 (US); inter Taco et Nibujon in ?charrascales-pinales? prope amnem, 4 December 1914, E. Ekman 3730 (S, holotype of Gesneria norlindii Urban); slopes of Sierra Azul, Quibijon, Baracoa, 400 m, Alain Lio- gier & M . Lopez F . 7318 (US); pine barrens, N. of El Yunque de Baracoa, 13 January 1960, Alan Liogier & J. Acuin 7626 (US); Baracoa in pinet. prope El Yunque (ad septentr. versus), 20 December 1914, E. Ekman 3564 ( S ) ; Baracoa, Lomas de Cuaba, edge of manacales, 17 January 1915, E. Ekman 4296 ( S , holotype o f Gesneria bracteosa Urban; K, S, isotypes); on top of El Yunque de Baracoa, 520 m, 2 January 1960, Alain Liogier & M . Lopez F. 7179 (US); northern slope of El Yunque de Baracoa, 450 m, 2 January 1960, Alain Liogier & M . Lopez F. 7179 bis (US); El Yunque (prope Baracoa) in cacum. montis. solo calcareo., 17-18 December 1914, E . Ek- man 3953 ( S , U S ) ; south of Monte Cristo, Baracoa, 7 Febru- ary 1992, J. Acuna & Diaz Barreto I7384 (SV, US 2 sheets); Baracoa, Lomas de Cuaba in fruticetis, 23 November 1914, E. Ekman 3611 (S); Cuchillas de Baracoa, 21 June 1860-1864, C. Wright sn (GH, holotype of Pentarhaphia duchartreoides C. U?right; GOET, isotype): 5 km S of Sabanilla, Via Azul, 350 m, 17 January 1960, Alain Liogier & J . Acuiia 7707 (US); 5 km S of Sabanilla, Via Azul, 17 January 1960, 390 m, Alain Liogier & J. Acuna 7710 (US); on the Via Azul between SMITHSONIAS COSTRIBUTIOA?S T O BOTANY Sabanilla and Cajababo, a t 5 km from Sabanilla, ca 500 m , 14 January 1956, C. Morton & Alain Liogier 9077 (US): banks of a rivulet, pinelands, km 7 S of Sabanilla, Via Azul, Bara- coa, 14 January 1956, Alain Liogier & C. Morton 5129 (IJ): Pinelands, km 7 south of Sabanilla, Via Azul, Baracoa, 14 January 1996, Alain Liogier & C. Morton 5140 (IJ); on the Via Azul between Sabanilla and Cajababo, a t km 7 from Sabanilla, ca 600 m , 14 January 1956, C. Morton & Alain Liogier 9089 (BM, US); on the Via Azul between Sabanilla and Cajababo, a t km 20 from Sabanilla, ca 800 m , 14 Janu- ary 1956, C. Morton & A h i n Liogier 9121 (BM, US); on the Via Azul between Sabanilla and Cajababo, a t km 22 from Sabanilla, ca 900 m , 12, 14 January 1956, C. Morton 8: A h i n Liogier 8992 (US): Sabanilla to Yamuri [sic] Arriba, 30 January-1 February 1911, J. Shafer 8420 (NY, US). DiscussIoN.-The holotype of Gesneria duchar- treoides, C . Wright sn, is deposited in the Gray Herbarium, based on information given by Hitch- cock (1934) concerning the location of type speci- mens of species names published by C. Wright and F. A. Sauvalle. Efforts to obtain information from Cuba concerning Wright collections extant in Cuba have been unsuccessful. From vegetative material it is very difficult to distinguish this species from Gesneria viridiflora; however, characters of the usually filiform calyx lobes and nearly cylindrical, verrucose corolla of Gesneria duchartreoides distinguish it from the Cuban material of Gesneria viridiflora with its apically thickened calyx lobes and campanulate, nonverrucose corolla. Section 5. Gesneria Conradia section Synanthera G . Don, Gen. Syst. 4:650, 1839. [Lectotype-species: Conradia humilis (Linnaeus) Martius ex G. Don.] Gesneria section Conradia (Martius) Fritsch in Engler & Prantl, Nat. Pflanzenfam. 4 (3b):184, 1894 [as to descrip- tion, not as to type], Plants acaulescent or stem erect, decumbent or pendent, to 0.5 m tall, branching from the base. Leaves oblanceolate to obovate or ovate, cuneate, membranous, plane, glossy or dull. Inflorescences exceeding the subtending leaves, either peduncle very long and pedicel very short, or peduncle very short and pedicel long; corollas tubular, red, orange, yellow, green, or white: sta- mens slightly exserted or included. Capsule forming a splash cup, costae prominent or obscure. TYPE-SPEcIES.-Gesneria humilis Linnaeus. DIscussIos.-Section Gesneria contains a seem- ingly diverse assemblage of plants, but they are combined into a somewhat natural group on the basis of habit, vegetative, inflorescence, and fruit characters. The relationships of this section to others, as well as its origin and position in a phylo- genetic sequence in the genus are unclear. Section Gesneria resembles section Physcophyllon in habit, corolla, and fruit characters and may be closely related, but differs in inflorescence and vegetative traits; however, an origin by reduction from some member or ancestor of section Pentarhaphia may be suggested. 23. Gesneria brachysepala Urban & Ekman Gesneria brachysepala Urban & Ekman in Urban, Ark. Bot. 2414 (4):34, 1932. Plants acaulescent or suffruticose: stems woody at the base, to 10 cm tall, 6-9 mm in diameter, bark reddish-brown, villous, lenticels obscure, resin not evident; unbranched, internodes very short, leaf scars prominent. Leaves alternate: petioles sulcate, 0.5-1.4 cm long, 2-4 mm wide, pilose with reddish or colorless trichomes; blades narrowly obovate, 5.6-1 3.9 cm long, 2.0-4.4 cm wide, subcoriaceous, base cuneate, margin crenate, subentire toward the base, apex rounded to acute, adaxial surface green, when young covered with broad-based trichomes, at maturity only bases remain to give a scabrous texture, abaxial surface green, with scattered appressed hairs especially along the prominent veins. Inflorescences 1- to 4 (-many)-flowered: pedun- cles terete, 9.1-20.4 cm long, 1 mm in diameter, red, glabrescent; bracts 2, linear, about 2 mm long, less than 1 mm wide, red; pedicels terete, 0.9-2.3 cm long, less than 1 mm in diameter, reddish, glabrescent; floral tube turbinate, ca 4 mm long, 3 mm wide, reddish, glabrescent; calyx lobes 5, connate for 1 mm at the base above ovary, erect to patent, with open aestivation, each lobe triangu- lar, 2.0-2.5 mm long, 1.5-2.5 mm wide, reddish, glabrescent on the outside, inside reddish to green, glandular, veins not prominent; corolla tubular, ventricose at the middle, narrowed to both ends, 2.6-3.7 cm long, 6 mm wide at the base and apex, NUMBER 29 91 Key to the Species of Section Gesneria 1. Pedicels more than four times as long as the peduncles; corolla orange-yellow to bright yellow, base and lobes of limb sometimes reddish: Puerto Rico.. ......................... 24. G. citrinn 1. Pedicels as long as the peduncles or less; corolla yellow-green, orange to red, or white. 2. Corolla red, more than 2.6 cm long; Haiti ............................................... 23. G. brachysepala 2. Corolla orange to yellow-green or white, less than 2.3 an long. 3. 3. 0.9-1.2 cm wide at red (fide Ekman), Corolla tube projected f horizontally from a nonoblique base, lobes of the limb erect, widest part of the tube at or near the middie; Puerto Rico ........ 26. G. pmciflara Corolla tube bent downward from an oblique base, widest part of the tube near the mouth, lobes of the limb patent or reflexed; Cuba and Haiti 25. G. humilis the middle, outside brilliant glabrous, inside yellow, gla- brous, limb very shortly 5-lobed, 5-8 mm wide, each lobe broadly obovate, reflexed, 1-2 mm long, 2-3 mm wide, dark red, margin ciliate; stamens 4, adnate to the base of corolla tube for less than 1 mm, filaments linear, exserted for 2 mm beyond mouth of corolla, curved, ca 3.5 cm long, 1 mm wide, yellow, glabrous, anthers narrowly rectangu- lar, 2-3 mm long, less than 1 mm wide, yellow, glandular, coherent in 2 pairs by their apices, staminode ca 6 mm long; ovary inferior, disc an- nular, puberulent with glandular trichomes, 2 mm long and 2 mm wide, reddish, style curved, 3.0-3.5 cm long, yellow, glandular, stigma clavate, papillate. Capsule broadly turbinate, becoming a splash cup, ca 4 mm long, 5 mm wide, gray at maturity, costae 5-10, obscure; seeds fusiform, less than 1 mm long, black. TYPE-COLLECTION.-LeS Roseaux at Debarcadere- Plingue, Haiti, E. Ekman HI0776 (S, holotype, Figure 53a; A, EHH, F, G, GH, IJ, K, NY, S, US, isotypes). DISTRIBUTION A D EcoLocY.-Gesneria brachysep- ala is endemic to the southern peninsula of Haiti, Departement du Sud (Figure 54) on steep shaded cliffs overhanging the sea, down to sea level. Ges- neria brachysepala has only been collected in October. This species would seem to be adapted for pollination by a hummingbird, but no known hummingbird species in Hispaniola has a bill long enough to reach the base of the corolla tube. Per- haps the nectar collects at the base of the ventri- cose middle and is thus accessible to shorter- billed birds. SPECIMENS ExAMINED.-HAITI. DEPARTEMENT DLI SLID: Massif de la Hotte, western group, Les Roseaux at Debarcadere-Plinguk [Plique], steep shaded cliffs, down to sea level, 4 October 1928, E . Ekman HI0776 (S, holotype of Gesneria brachysepala Urban & Ekman; A, EHH, F, G, GH, IJ, K , NY, S, US, isotypes). DrscussIoN.-Gesneria brachysepala has been col- lected only once. T . Talpey and I sought this rare species at the type locality in July 1970 but failed to find it. Residents in the area reported that the cliff face, where presumably Gesneria brachysepala grew, was destroyed by the effects of a hurricane a number of years ago. Plants were not located on nearby cliffs. Gesneria brachysepala resembles other members of this section in its habit, inflorescence structure, and splash-cup fruit. The species may be distin- guished from others, however, in its red tubular corolla up to 3.7 cm long. 24. Gesneria citrina Urban Gesneria citrina Urban, Symb. Ant. 1:477, 1900.-Britt. & Wils., Sci. Surv. P.R. & V.I. 6:205, 1925. Subshrubs: stems woody at the base, pendent or decumbent, to 1 m long, 2-5 mm thick, bark of young stems reddish to brown, becoming brown to gray, glabrescent, apex slightly resinous, lenticels sometimes erumpent, elongated, white to brown; pith reddish; branches from the stem base, rarely in upper leaf axils, internodes 0.1-1.0 cm long, leaf scars swollen. Leaves alternate, petiolate or sessile, persistent: petioles terete to subsulcate, 0-1.0 cm long, 1-2 mm in diameter, red to gray-brown, sparsely pubescent with glandular or eglandular trichomes or gla- brous; blades obovate or spathulate, 1.2-5.9 cm long, 0.5-2.9 cm wide, coriaceous, plane, base cuneate, margin ciliate to glabrescent, serrate to dentate toward apex, apex acute to obtuse, adaxial surface dark green, glabrous, glossy, abaxial sur- 92 SMITHSONIAN CONTRIBUTIONS T O BOTANY FIGURE 53.-Type specimens: a, holotype of Gesneria brachysepala Urban & Ekman, Ekman HI0776 (S); b, lectotype of G. citrina Urban, Sintensis 6590 (S). face lighter green, becoming white when dry, glabrous, occasionally with a few appressed tri- chomes along the midvein, glossy, veins sometimes reddish-brown, prominent, dried leaves with ex- foliating epidermis. Inflorescences 1- to 4 (-co)-flowered: peduncles very short, 1-2 mm long, flattened to terete, about 1 mm wide, red, glandular-pubescent, occasionally resinous; bracts 2-4, linear, 1-3 mm long, less than 1 mm wide, red, resinous; pedicels terete, arcuate, 2.8-6.3 cm long, about 1 mm in diameter, red, sparsely pilose; floral tube shortly turbinate, 2 mm long, 2-3 mm in diameter at anthesis, green to red, sparsely pilose to glabrous, resinous; calyx lobes 5 , connate for 1-2 mm at base, each narrowly triangular, 2-3 mm long, 1.0-1.5 mm wide at the base, apex acute to acuminate, reddish or green, sparsely pilose, veins 3, prominent, inner surface with short glandular trichomes; corolla tube, curved, nearly cylindrical, 1.6-2.1 cm long, 3 mm wide at the sometimes reddish base, widening to 7 mm at the slightly (Figure 16i) ventricose middle, then narrower toward the oblique mouth where 5 mm wide, outside orange-yellow to bright yellow, densely puberulent, inside lighter yellow, glabrous, limb 5-lobed, erect, each lobe semiorbiculate, glabrous, yellow or sometimes reddish, margin sub- entire, often ciliate, upper lobes 1 mm long, 2 mm NUMBER 29 93 HISPA NIO 1 A , IOOkm I FIGURE 54.-Distribution of Gesneria section Gesneria and Section Physcophyllon in Hispaniola. (G. (Physcophyllon) barahonensis = open triangles; G. (Gesneria) brachysepala = solid circles; G. (Physcophyllon) christii = solid squares; G . (Gesneria) hurnilis = open circles; G. (Physco- phyllon) hybocarpa = stars; G. (Physcophy Elon) pedicellaris = open squares; G. (Physcophy lh ) reticdata = solid triangles.) wide, lateral lobes 3 mm long, 2.5 mm wide, basal lobe 2.5 mm long, 2.5 mm wide; stamens 4, adnate to the base of the corolla tube for less than 1 mm, not exserted, filaments linear, straight, 1.6-1.9 cm long, less than 1 mm in diameter, yellow or white, glabrous, anthers rectangular, 1.5 mm long, 1 mm wide, yellow to red on back, coherent in two pairs by their apices, pollen grains isopolar, with size small to medium (18.8-25.6 pm long at the polar axis, 12.3-16.0 pm wide at the equatorial axis), with a nearly circular amb, tricolpate, colpi 20 pm long, less than 1 pm wide, apocolpia rounded to acute, prolate, sexine reticulate, heterobrochate, lumina 0.1-1 .O pm across, larger near the equatorial axis, muri about 0.5 pm across; ovary inferior, disc annular, 1.5 mm long, green, puberulent with glandular trichomes, style straight except curved at apex, to 2.3 cm long, white, glabrous, stigma clavate, papillate. Capsule broadly turbinate, becoming a splash cup, 3-5 mm long, ca 6 mm in diameter, gray- brown, glabrescent, glossy, costae 5-10, prominent; seeds narrowly fusiform, less than 1 mm long, tawny. TYPE-CoLLEmI0N.-Prope Utuado, Puerto Rico, P. Sintenis 6590 (S, lectotype, Figure 53b; BM, BP, BR, C, E, F, G, GH, GOET, K, L, LD, M, MO, NY, P, UCWI, US, W, Z, isolectotypes). CHROMOSOME NUMBER.-n = 14 (Lee, 1966). DISTRIBUTION A D ECOLOGY.-Known from lime- stone cliffs in the hills of north-central Puerto Rico (Figure 55) at about 200-325 m elevation, Ges- neria citrina has been collected in flower during the months of February, March, May, August to November in the field. In the greenhouse it flowers in May and October. The corolla tube appears to be adapted for pollination by hummingbirds (Chlorostilbon maugaeus or Orthorhynchus cristatus). November 1913, F . Stevens & W . Hess 5163 (NY); Lares, 325 m, 24 October 1943, F . Sargent 3235 (US); near the Arecibo Ionospheric Observatory on P. R. Hwy 625 about 15 km S of Arecibo, 27 August 1970, L . Skog XC J . Skog 1716 (BH, E, K, US); Esperanza, v8 mi along path toward the left just before the entrance to Cornell Radar in Esperanza south of Arecibo, ca 800 ft, 12 February 1965, T . Talpey 28 (BH, US); Arecibo radar station, 27 February 1965, T. Talpey 41 (BH, US); cliffs, Arecibo, 31 May 1935, F . Sargent 703 (US); base of cliff between Arecibo and Utuado, 8 February 1924, N . Britton & E. Britton 7976 (GH, NY, PH, US); prope Utuado in sylva primaeva montis Cerro Hueco ad Cayuco, 25 March 1887, P. Sintenis 6590 ( S , lectotype of Gesneria citrina Urban: BM, BP, BR, C, E, F 2 sheets, G 5 sheets, G H GOET, K, L, LD, M, MO, NY, P, UCWI, US, W, Z, isolectotypes); Barce. loneta, S of Florida, 250 m, 23 August 1963, H . McKee 10616 (K, P, US); near Florida on route 140, 27 September 1967, R. Wagner 1233 (A); Rio Piedras, Ciales, 17 August SPECIhlENS ExAMINED.-PUERTO RICO: Quebradillas, 22 94 SMITHSONIAN CONTRIBUTIONS T O BOTANY b SO km PUERTO RlCO 4 FIGURE 55.-Distribution of Gesneria section Gesneria and section Duchartrea in Puerto Rico. (G. (Gesneria) citrina = solid circles; G. (Gesneria) paucipora = solid triangles; G. (Duchartrea) viridipora subsp. sintenisii = solid squares.) 1903, J . Johnston 974 (SY, US), LOCALITY UNKKOWN: Rio Arriba, Forest Reserve, I . Velez I606 (NY). CULTIVATED: Cor- nell University, G-888, 5 October 1969, L. Skog 1408 (BH), 23 May 1966, 111. Stone 130 (BH). DIscussIoN.-Gesneria citrina is distinct from all other species in this section in its short peduncles, 1-2 mm long; peduncles of the remainder of the species are more than 2 cm long. I ts subshrubby habit with long pendent or decumbent branches in combination with yellow tubular corollas make Gesneria citrina distinguishable from all other known species of Gesneria. From the many duplicates of the presumed de- stroyed holotype, Sintenis 6590 at Berlin, a sheet at Stockholm has been chosen as lectotype. None of the isotypes found in many herbaria display flowers or is annotated in Urban?s hand, but the specimen at Stockholm is typical of those examined. 25. Gesneria humilis Linnaeus Gesneria h i m i l i s Linnaeus, Sp. P1. ed. 1, 612, 1?53.-Plum., PI. Am, 125, pl. 133: fig. 2, 1757.-Lam., Encyl. Meth. Bot. 2:?02, 1788 [as to the type, excluding synonyms of Sloane and Ray].-Lunan, Hort. Jam. 1:322, 1814.-Poir. in Lam., Illustr. 3: PI. 536: fig. 2, 1819.-Spreng., Syst. Veg. ed. 16. 2:839, 1825.--Urb., Symb. Ant. 2:377, 1901. Conradia kumi l i s (Linnaeus) Martius ex G Don, Gen. Syst. 43650, 1838.-DC., Prodr. 7:526, 1839.-Rich. in Sag., Hist. Fis., Pol. Nat. Cuba 11:71, 1850.-Hanst., Linnaea 26: pl. 1: fig. 34, 1854.-Griseb., Mem. Amer. Acad. Arts, n.s. 8:526, 1862; F1. Brit. W. Ind. 461, 1862; Cat. P1. Cub. 200, 1866. Pentarkapkia humilis (Linnaeus) Hanstein, Linnaea 34:294, 1865. Conradia celsioides Grisebach, Cat. P1. Cub. 200, 1866. [Type- collection: Vinales, Cuba, C. Wright 3077 (GOET, holo- type: BM, G, GH, K, MO, P, S, TY, isotypes).] Pentarhapkia celsioides (Grisebach) Gomez de la Maza, Anales SOC. Esp. Hist. Nat. 23:279, 1894. Gesneria acurninata Urban, Symb. Ant. 1:479, January 1900.- Morton in Leon & Alain, F1. de Cuba 4:463, 1957. [Type- collection: Arroyo de Pedro, Cuba, H . Eggers 4892 (GOET, lectotype: C, P, Z, isolectotypes).] Gesneria incisa Urban, Symb. Ant. 1:479, January 1900. [Type-collection: In rupibus fluminum Cubae, E . Poepp ig sn (TV, lectotype; BP, BR, MO, isolectotypes).] Conradia p t i n i l a sensu Millspaugh, Publ. Field Columbian Mus. Bot. Ser. 1:434, August 1900 [non Conradia p u n i l a (Swartz) Martius (1829 et 1832).] Gesnevia celsioides (Grisebach) Urban, Symb. Ant. 2:377, 1901.-Morton in Leon & Alain, F1. de Cuba 4:463, fig. 201, 1957. Subshrubs: stems woody, erect or occasionally pendent or decumbent, to 0.5 m tall, bark greenish- gray, smooth, rarely resinous, lenticels obscure; branching at the base or rarely above, new growth green, pilose to glabrous, internodes 0.1-2.0 cm long, leaf scars prominent. Leaves alternate and spiraled, petiolate or sessile: petioles flattened to sulcate, 0-1.3 cm long, 1-3 mm wide, light or dark green or reddish- brown, sparsely glabrous to pubescent-glandular; NUMBER 29 95 blades narrowly oblanceolate to narrowly obovate or ovate, 1.3-18.0 cm long, 0.6-4.3 cm wide, mem- branous, plane, base cuneate, margin serrate to crenate or lobulate, subentire toward base, sparsely ciliate to glabrous, apex acute to acuminate, ad- axial surface dark green, sparsely pilose near the base, trichomes appressed, sometimes with broad bases, glossy to dull, abaxial surface lighter green, sparsely pilose to glabrescent on the prominent veins. Inflorescences 1- to 3 (-m)-flowered: peduncles terete, 0.4-17.8 cm long, elongating in fruit, 1.5- 2.0 mm in diameter, green to reddish, usually glabrous; bracts 2, linear to lanceolate, 0.1-1.0 cm long, 1-3 mm wide, green, glabrous, base cuneate, apex acute; pedicels terete, 2.2 cm long, less than 1 mm in diameter, green to reddish, glabrous; floral tube turbinate to nearly spherical, 1-3 mm long, 2-3 mm in diameter, dark green or reddish, glabrous; calyx lobes 5 (-7), erect, connate for 1-2 mm above ovary, with open aestivation, each lobe lanceolate or triangular, 1.0-4.5 cm long, 1-3 mm wide at the base, apex acute to acuminate, both sides green or reddish and glabrous, veins 3, not prominent, but converging at apex; corolla tube narrowly campanulate, 0.6-2.0 cm long, base oblique, 2-4 mm wide, gradually (Figure 161) widening to ventricose on the lower side, slightly constricted at the mouth, 3-8 mm wide, greenish- white to yellow, glabrous outside and inside; limb 5- or rarely 6-lobed, 4-8 mm wide, each lobe patent to reflexed, semiorbiculate, green to white, rarely pinkish, 1-5 mm long, 3-6 mm wide, upper lobes subentire to erose, glandular, lateral lobes erose to subentire, basal lobe subentire; stamens (3-)4 (-5), adnate to the base of corolla tube for less than 1 mm, not exserted, filaments linear, geniculate, 0.4- 1.3 mm long, green to yellow, glabrous, anthers triangular to globose or oblong, ca 1 mm long, 0.5-1.0 mm wide, green, seldom coherent, pollen grains often distorted, isopolar, size small (15.1 pm long at the polar axis, 8.5 pm wide at the equa- torial axis), tricolpate, colpi 11.5 pm long, less than 1 pm wide, apocolpia acute, prolate, sexine reticu- late, nearly homobrochate, lumina 0.25-0.5 pm across, muri ca 0.5 pm wide (Figure 17h), staminode 1 or rarely 2, ca 2 mm long, lacking fertile anther; ovary inferior, disc an undulating collar, 1 mm tall, white, pubescent, style linear, curved, 0.5-1.4 cm long, about 0.5 mm in diameter, green, gla- brous, stigma slightly enlarged, stomatomorphic, 1-3 mm across, occasionally adherent to the anthers. Capsule turbinate to nearly spherical, forming a splash cup with the calyx lobes, 3-7 mrn long, 4-8 mm in diameter, gray-brown, glabrous, costae 10, usually prominent; seeds rhombic to broadly fusiform, slightly twisted, ca 0.5 mm long, 0.2 mm wide, brownish (Figures 19j, 20b). TYPE.-Plumier, Nov. P1. Am. Gen. 27, fig. 9, 1703. The type figure was only part of the original drawing made by Plumier shown in Figure 57. DISTRIBUTION A D EcoLoGY.-Gesneria humilis is known in Cuba (Figure 56) from the Provinces of Pinar del Rio, La Habana (including the Isle of Pines), Las Villas, and Oriente; and in Haiti (Fig- ure 54) from the Departement de l?Ouest. Plants grow in dense, damp limestone woods near streams or on rocks in streams. Gesneria humilis has been collected in the wild with flowers and fruits in all months of the year, but with fewer collections in late spring; greenhouse plants flower with the same regularity. T h e pollinators of this species have not been determined, but the plants may be visited by hummingbirds. LOCAL NAME.-?Clavellina? fide L u n n 21 7 (NY). SPECIMENS EXAMINED.-cUBA. PROVINCE OF PINAR DEL RIO: Jovero, Guanes, 19 January 1948, J. Acuna 14940 (SV, US); San Julian, in pineland savannas at the headwaters of Rio Verde, 6 June 1920, E . Ekman 11134 (S); Sierra de Cobra, on Guane Road, 9, 11 September 1910, N. Britton, E . Britton & C . Gager 7216 (NY), 7220 (NY, US); Surgidero, Rio San Vincente, Vinales, August, 1954, J. Acufia 8c Maza 19352 (SV, US); San Vicente, Viiiales, 2 February 1951, J. Acuna & J . Roig sn (US); vicinity of San Vicente, near ViAales, near sea level, 5 February 1956, C. Morton 9927 (US); base of Sierra del Ruisenor, Vinales, 21-22 May 1955, Alain Liogier 4322 (GH, NY); Vinales, Rio Pan de Azucar, 9 October 1955, Alain Liogier 4434 (US); Pan de Azucar, 20-100 m, 5 Febru- ary 1956, C . Morton 9918 (US); Banos San Vicente, 12-16 September 1910, N. Britton, E . Britton & C . Gager 7358 (NY), 7359 (NY); Vinales, Sierra de la Guasaca, 10 March 1924, E . Ekman 18683 (S); Vinales, 11 December 1930, E. Kil l ip 13579 (US); San Vicente, April 1937, Bro. Leon 16808 (US); Cueva de San Vicente, San Vicente, Vinales, 2-3 Sep- tember 1937, Bro. Leon 16960 (GH, US); El Guao, Vinales, July 1939, Bro. Leon 19045 (US); El Valle de Ancbn, near San Vicente, 4 February 1956, C . Morton 9773 (US); base of Sierra del Ruisenor, Finca El Ancon, near Vinales, ca 150 ft, 17 March 1957, G. Proctor 16357 (IJ); on cliff, Vinales, 1860- 1864, C. Wright 3077 (GOET, holotype of Conradia celsioi- des Grisebach; BM, G 2 sheets, GH, K , MO, P, S, W, iso- types); Arroyo del Sumidero, 7, 9 August 1912, J. Shafer & 96 SMITHSONIAN CONTRIBUTIONS T O BOTANY FIGURE 56.-Distribution of Gesneria section Gesneria and section Duchartrea in Cuba. (G. (Ges- neria) humilis = open circles; G. (Duchartrea) viridiflora subsp. uiridiflora = solid circles.) Bro. Leon I3601 (F, XY, P H , US); in mountains near El Guama, 6 March 1900, W . Palmer & J . Riley 153 (US); vicin- ity of Pinar del Rio .Arroyo, 3 March 1911, A?. Britton, E. Britton & J . Cowell 9723 ( S Y ) ; La Cajalbana, 1 October 1949, J . A c u n a & Alain Liogier I5686 (SV, US); Loma de la Cajzilbana, 1 October 1949, J . AcuNa & Alain Liogier sn (IJ 2 sheets, US); in water of a waterfall, Cajailbana Mt., La Palma, 10 October 1949, Alain Liogier 1185 (GH, US); in a stream, Cajailbana pinelands, La Palma, 7 December 1955, Alain Liogier 4491 (US); Pinar del Cajzilbana, in savannas a t the foot of mountains, on the edge of a branch of Rio Puercos, 28 August 1923, E. Ekman 17311 ( S ) ; banks of the river, San Diego de 10s Banos, 19 August 1914, Bro. Leon 4410 ( S Y , US 2 sheets); in mountains north of San Diego de 10s Banios, 11 April 1900, TV. Palmer Bc J . Riley 520 ( S Y , US); rocky stream bed, San Diego de Los Banos, 31 August-3 Septem- ber 1910, N . Britton, F. Earle 8: C. Gager 6740 ( S Y 2 sheets); rocky stream bed, San Diego de Los Banos, 31 August-3 September 1910, M. Britton, F. Earle k C. Gager 6889 ( S Y , US); Bahia Honda, 31 March 1946, J . X o i g , J . A c u n a R: Naranjo sn (US); Rio Taco-Taco, Rangel, August 1927, J . A c u n a sn (US); mountains above Taco-Taco, C. Baker 3832 (XY); source of Rio Taco-Taco, Sierra de Los Organos, 400-500 m , 18 Sovember 1941, C. Morton 4283 (BM, F, G , GH, hfO, SY, S, U, US); Rangel, Arroyo de la Plata, 23, 24 December 1951, Alain Liogier Bc E. Kil l ip 2010 (US); vicinity of Guane, falls, Rio Portales, 3 hfarch 1911, X. Britton, E . Britton 8i J . Cowell 9765 (NY); Rio Portales, 26 December 1911, J . Shnfer 11171 (F, G H , MO, S Y , U, US); Bahia Honda to El Rosario, crossing of San Miguel, 29 January 1912, J . Shafer I2000 (I(, S Y 2 sheets, U, US); Bahia Honda to El Rosario, 29 January 1912, J . Shafer 12002 (MO, SY 2 sheets); Cascada de Saroa, Candelaria, 5 March 1951, J . Acuna s n (SY); Rio Sfananteales north of Candelaria, 10 February 1916, A?. Britton, P. Wilson & Bro. Leon 14127 (SY); San Pedro del Caimito to San Jose de Sagua, 27 January 1912, J . Shafer 11956 (MO, NY, US). PROVINCE OF LA HABANA, ISLE OF PINES: vicinity of Los Indios, 13 February 1916, A?. Britton, E . Britton R: P. Wilson 14241 (GH, hIO, NY, S): rocky ford south of Santa Barbara, 11 March 1953, E. Killip 43088 (F, XY, US); between San Francisco de las Piedras and Cerro La Canada, 27 January 1955, E. Killip 44616 (US); dense woods along river, Razlag property along Rio Ma1 Pais, 25 December 1955, E. Killip 45270 (US); in savannas a t Loma Daquilla, 3 December 1920, E. Ekntan 12496 ( S ) ; Santa Fe, cerca del Rio l?ellegas, 15 March 1923, M . Caluino R: E. Manzteli 7988 (SV, US); Santa Fe, 15 March 1953, M. Caluino in Leon 22385 (US); bank of stream, Santa Fe, 8 March 1953, E . Killip 43050 (US); dense woods along stream, Mrs. Jones? Jungle, 7 March 1957, E. Killip 45830 (US); ?Isla De Pinos,? June, J . Blain 50 (F). PROVINCE OF LA HABASA: asphalt mines, Santiago de las Vegas, 18 Septem- ber 1904, H . van Hermann 42 (BM, F, NY); on rocks in rapids of stream, Bajucal, 5 December 1904, H . uan Hermann 332 (F, SY); near asphalt mines above Bajucal, 18 Septem- ber 1904, H . 7 m n Hermann 1648 (NY); Loma Coca near Campo Florida, at Rio Quezada, 21 September 1921, E. Ekrnnn 13226 ( S ) ; banks of the river near Cumbre Hermosa, 3 January 1912, P. Wilson k Bro. Leon 2817 (SY); on rocks in stream, Cumbre Hermosa, 3 January 1912, P. Wilson & Bro. Leon 11617 i S Y ) ; ?Havanna,? G. Don sn (GOET); ?Havana,? 1833, X. d e / a Sagra 647 (G-DC). PROVINCE OF LAS VILLAS (formerly Santa Clara): San Blas, 17 March 1929, L. Bailey 12f33 (BH); above San Blas (La Sierra), 7 April 1928, J . Jack 5951 (.A, LS); Loma Ventana, San Blas, La Sierra, 1400 ft, 22 July 1930, J . Jack 8035 ( S , US): Loma l?entana, San Blas-Las Vegas, La Sierra, 1400 ft, 24 July 1930, J . Jack 8067 (.A, S, US); San Blas, 300-600 m, 9-10 Sovember 1941, C . Morton 4110 (Bar, US); Mina Carlota, SE of Cumanaya- gua, Sierra tle San Juan, 300-400 m, August 1940, R. Howard 4480 (A); Arroyo Savarro, hfina Carlota, southeast of Cu- manayagua, Sierra de San Juan , 300-400 m , 7 July 1941, R. Howard 5716 [PI. ex. Crayanae 11821 (BH, BM, BR, C, COL, CU, E, G , G H , I J , I(, L, LD, hfO, NA, KY, NYS, P H , S, U, US, USF); Arroyo Savarro, hlina Carlota, in Sierra de San Juan southeast of Cumanayagua, ca 1000 ft, 22 March 1957, G. Prortor 16403 (IJ); hlina Carlota, southeast of Cumana- yagua, Sierra tie Sail Juan , 300-400 m , 21-23 March 1938, H . Senn 3?2 (CH, US); Hanabanilla Falls, 1-20 July 1950, R. Hou?ard, W . Briggs, P. Kamb, I . Lane & R. Ritland 148 (A); NUMBER 29 97 Hanabanilla Falls, Trinidad Mountains, 23 February 1956, C. Morton 10446 (US); Hoyo de Manicaragua, 26-28 Febru- ary 1910, N . Britton, E. Britton & P. Wilson 4695 (F, NY 2 sheets); El Porvenir, Trinidad Mountains, 650-750 m, 9 March 1910, N . Britton & P. Wilson 5274 (NY); western slopes of Mt. Naranjal above San Blas, Trinidad Mountains, 28 July 1936, L . Smith, A . Hodgdon & F . Gonzales 3248 (F, GH, MO, NY, S , US); Buenos Aires, Trinidad Mountains, 21 February 1956, C. Morton 10309 (US); Lomas de Banao, February 1920, A . Luna 217 (NY); Madrigal near Sancti Spiritus, 27 August 1909, Bro. Leon 948 (NY); vicinity of Sancti Spiritus, 15-24 February 1912, J . Shafer 12165 (F, MO, NY, US); Pitajones to Ciegas de Ponciano, 29 February 1912, J. Shafer 12254 (NY, U, US). PROVINCE OF ORIENTE: Rayate, at margin Arroyos Vivano, 18 July 1914, E. Ekman 1993 (K, S ) ; Bayate in saxis in Arr. Bibano, 5 May 1919, E. Ekman 9615 ( S ) ; Puerto Boniato, Santiago de Cuba, August 1949, Bro. Clemente 6998 (GH, US): Sierra de Nipe, El Taller in saxis in flumine Rio Piloto, 3 November 1914, E. Ekman 3324 (G, K , S ) ; Manantial, Sta. Fe, N. de Guantanamo, 7 December 1917, Bros. Hioram & Btiste 1380 (NY, US); Guan- tanamo, Monte Libanon, San Fernandez ad amnem in ?manacales,? ca 700 m, 24 December 1919, E. Ekman 10243 ( S ) ; Arroyo Henequen, foot of El Yunque, 1 December 1910, J . Shafer 7695 (NY); Veriles, sur de Monte Cristo, 7 Febru- ary 1952, J . Acuna & Diaz 17453 (US); Arroyo de Pedro, 500 m, March 1889, H . Eggers 4892 (GOET, lectotype of Ges- neria acuminata Urban; C, P, Z, isolectotypes); ?in Cuba Orientali,? 1856-1857, C. Wright 355 (BR, G, GH, GOET, K, MO, NY, PH); prope villam Monte Verde dictam, Jan- uary-July 1859, C. Wright 355 (GH, GOET, K); ?in Cuba Orientali,? 1860, C. Wright 355 (K, W); in Rio Santa Cruz and others, 12 October 1860-1864, C. Wright 355 (BM); ?in Cuba Orientali,? 1860-1864, C. Wright 355 (BM, BR, G, GOET, P). LOCALITY UNKNOWN: window ravine, 2 March 1920, J . Bijhouwer 428 (WAG); Santa Catalina, Sierras of Western Cuba, January-February 1907, H . Caldwell & C. Baker 7029 (F); ?In rupibus fluminum Cubae,? E. Poeppig sn (W, lectotype of Gesneria incisa Urban; BP, BR, MO, isolectotypes); ?Flos. Cubens.-ad torrentio.,? July [18]23, E. Poeppig sn (P, S); ?Cuba,? 1822, E . Poeppig sn (K); ?Cuba,? R. de la Sagra sn (W); ?Obre 833, Legit R. de la Sagra,? [J. M.] Valenzuela sn (P); ?Cuba occ.,? 1863, C. Wright 647 ( S ) . HAITI. DEPARTEMENT DE L?OUEST: plants collected from the base of a volcanic rock cliff at Fond de Baudin along Rivikre Toreau [R. Gauche?] about 2.5 mi NE of Trouin, 18O23? N lat. 72038? W long., 1100 ft, 22 July 1970, L . Skog, T . Talpey & D . Pfister 1610 (toPotype: BH, MO, US). DIscussIoN.-Gesneria humilis was first discov- ered in Haiti by Plumier who described the species as Gesnera humilis, flore flavescente in 1703. No collections from Haiti, however, were known until its rediscovery in 1970 (Skog and Talpey, 1971). This species has been widely collected in Cuba, where it is the most common species of Gesneria, according to Morton (195ib). Specimens from both islands are very similar except in pollen. Pollen from plants collected in Haiti appears distorted and collapsed under scan- ning electron microscopy (Figure 1 S f ) ; samples from Cuban collections appear to be normal (Fig- ure l ih) . The Haitian plants, however, produce viable seed. One population at Viiiales, Cuba, has been separated as a distinct species, Gesneria celsioides (Grisebach) Urban. The characters, however, that distinguish this species may be the result of FIGURE 57.-Original manuscript drawing by Plumier of Gesnera humilis, pore pavescente from the Bibliotheque Centrale of the Museum National d?Histoire Naturelle at Paris. (See Figure 58 for representative specimen of Gesneria humilis from Haiti.) 98 SWITHSOSIAS CONTRIBUTIONS T O BOTANY FIGURE 58.-Representative specimen of Gesneria hurnilis Linnaeus from Haiti, Skog, Tal$ey & Pfister 1610 (BH). ecological conditions. Other populations dwelling in or near streams appear distinctive in being much reduced, perhaps because they are often in- undated. Neither of these populations is here con- sidered as a taxon separate from the typical Gesneria humilis. Both Gesneria acuininata and G. incisa were based by Urban on more than one collection. It is presumed that the original syntypes were at Berlin and destroyed, thus, it is necessary to select lecto- types of these species. Urban based Gesneria acuminata on Wright 355, Valenzuela 10, Eggers 4892 and 4892b. T h e Valenzuela specimen has not been located in any herbarium, nor has the second Eggers number been found. Wright specimens bearing the same number were not necessarily collected at the same time nor at the same site; some specimens with the number Wright 355 bear the date 1856-1857 and others the date 1860-1864. Often two speci- mens with different dates but the same number are mounted on the same sheet. It is impossible to determine which specimen corresponds to the ap- propriate date. Those dated 1856-1857 give as locality only ?Cuba orientali.? The others are- lo- cated as ?prope villam Monte Verde dictam, Cuba orientali.? In still other herbaria, collection dates are either July 7, 1859, Jan-Jul 1859 or Oct 12, or labeled as coming from ?in Rio Santa Cruz and others.? It would be difficult to select one of the Wright specimens to be the lectotype due to the disparate dates and localities. Only a few are labeled in Urban?s handwriting. I suspect, however, that the Wright 355 Urban listed as a syntype was a 1856-1857 collection, for if the specimen had been collected in 1860-1864 it would have also been labeled with a definite locality. Thus by elimination I have selected Eggers 4892 (GOET) as the lectotype for Gesneria acuminata. I t is labeled in Urban?s handwriting and has a definite collection locality and date. The specimen has both flowers and fruiting material present. Gesneria incisa was based on specimens collected in Cuba by E. Poeppig and R. de la Sagra. If Urban had specimens from these collections at Berlin, they are now not extant. Other specimens of these collectors can be found in many herbaria, but none has been found to have any annotation by Urban. I have selected a Poeppig specimen in the herbarium at Vienna to be the lectotype for Gesneria incisa. It is a representative specimen with both flowers and fruits. Many authors, including de Candolle and Urban, have cited Martius as the author who made the combination Conradia humilis, since it was listed in 1829 as one of the species of Gesneria Linnaeus that Martius proposed to include in his new genus. T h e combinations of the Gesneria species in Conradia were made in the index in 1832, except for Gesneria hunzilis Linnaeus and G. ventricosa Swartz. Apparently through an over- sight these species were not included in Martius? index. G. Don finally made the transfers in 1838. 26. Gesneria pauciflora Urban Gesneria pamiporn Urban, Symb. Ant. 1:478, 1900.-Urb. S p b . Ant. 2:381, 1901; 4:57l, 1911.--N. Britt. & P. Wils., Sci. Surv. P. R. & V. I. 6:205, 1929. NUMBER 29 99 Plants acaulescent or suffruticose: stems woody at the base, erect or decumbent, to 3 dm tall, to 8 mm thick, bark smooth, gray-brown, glabrescent, lenticels obscure; branches few from the base of the stem, deep red, pilose with appressed hairs, internodes 1 cm long or more, leaf scars slightly enlarged. Leaves alternate: petioles terete or flattened, 2-7 mm long, 1-2 mm wide, green, pilose; blade narrowly obtrullate, 2.8-9.2 cm long, 0.9-2.3 cm wide, membranous, plane, base cuneate, margin subentire toward the base, serrate to sublobate above, ciliate when young, becoming glabrescent, apex acuminate, adaxial surface dark green, glabrescent, glossy, abaxial surface lighter green, pilose along the prominent veins. Inflorescences 1- to few-flowered: peduncles terete, 6.1-15.3 cm long, 1-2 mm in diameter, slightly curved, reddish-brown, pilose; bracts 2, linear, ca 1 cm long, 0.5 mm wide, green, pilose; pedicels terete, 1-2 cm long, 1 mm in diameter, reddish-brown, pilose to glabrescent; floral tube shortly turbinate, ca 2 mm long, 3 mm wide at anthesis, green, pilose; calyx lobes 5, connate for about 1 mm at base, erect, with valvate or open aestivation, each lobe narrowly triangular, 3-4 mm long, 1-2 mm wide at the base, green, reddish toward the acuminate apex, margin entire, outside pilose to glabrescent, inside glabrous, veins 3, not prominent; corolla (Figure 16k) tubular, curved, 2.0-2.3 cm long, 4 mm wide at the base, narrowing to 3 mm wide, then widening at the ventricose middle to 5 mm and narrowing at the mouth to 4 mm, outside yellow-orange, densely pilose with appressed hairs, inside glabrous, limb 5-lobed, 6 mm wide, each lobe erect, broadly elliptic, yellow with an entire, ciliate margin, upper lobes 1 mm long, 3 mm wide, lateral lobes 2.5 mm long, 2 mm wide, basal lobe 4 mm long, 3 mm wide, mouth lined with glandular trichomes; stamens 4, adnate to the base of the corolla tube for 1.5 mm, not exserted, but extending to within 2-4 mm of the mouth, filaments linear from a broad base, 1.5-1.8 cm long, 1 mm wide at the base, yellow, pilose only near the base, glabrous above, anthers broadly ovate, less than 1 mm long, yellow, co- herent in 2 pairs by their apices, becoming free, pollen grains isopolar, size small (19.0 ym long at the polar axis, 11.0-11.5 pm wide at the equatorial axis), amb nearly circular, tricolpate, colpi 17.0 ym FIGURE 59.-Lectotype of Gesneria puzlcifloru Urban, Sintenis 327 (GOET). long, apocolpia truncate, sexine reticulate, homo- brochate, lumina 0.5 pm wide, muri 0.25 pm across, (Figure 18e), staminode 7 mm long, adnate to corolla for 4 mm; ovary inferior, disc a 5-angled thick ring, yellow-green, pilose, style linear, slightly curved, 1.4-1.9 cm long, yellow, glabrous, becom- ing puberulous near the apex, stigma stomato- morphic, papillate. Capsule becoming a splash cup, ca 4 mm long, 4 mm wide, gray-brown, glabrescent, costae 5-10, not prominent; seeds broadly fusiform, twisted, less than 1 mm long, ca 0.5 mm wide, dark reddish-brown. TYPE-COLLECTION.-Maricao, Puerto Rico, P. Sintenis 327 (GOET, lectotype, Figure 59; BM, BP, G, GH, K, LD, M, P, S, US, isolectotypes). 100 SMITHSOSIAS C O S T R I B U T I O K S TO B O T A S Y CHROMOSOME NUMBER.-n = 14 (Lee, 1966). DISTRIBUTION A D EcoLow.-Gesneria pauciflora is endemic to the mountains of western Puerto Rico (Figure 55) in wet forests at stream sides at elevations from 500-720 m. Gesneria paucipol-a has been collected in flower in all months except March, July, and September; in the greenhouse flowers may appear every month. Hummingbird pollination is suspected in this species. Sel-icotes holosericeus with a bill averaging 21 mm long may be adapted to feed from Gesnel-ia pauciflol-a. SPEClMEr\.S ExAMI~\.ED.-PC?ERTO RICO: River valley, Rio de Maricao, 600-720 m, 14 February 1915, A?. Britton 8: J . Corcell 4192 ( S Y ) ; Indiera Fria, near Maricao, rocky arroyo, 500 m, 19-22 February 1915, A?. Brittoti, J . Cowell 8: S. B r m n 4546 (F, MO, NY, US); on rocks in wooded valley, Rio de Maricao, 500-600 m, 2 April 1913, N . Britton, F. Stevens 8i TV. Hess 2442 (F 2 sheets, GH, MO, KY 2 sheets, US 2 sheets); Maricao in rupibus ad rivulum in ?Indiera-Fria,? 3 December 1884, P . Sintenis 327 (GOET, lectotype; BM, BP, G 2 sheets, GH, K , LD, M, P, S, US, isolectotypes); along trail near Rio Maricao about 1 km S of the fish hatchery a t Maricao, 550 m, 13 June 1970, L . Skog 1539 (BH), 16 June 1970, L. Skog 1542, 1544 (BH), 26 August 1970, L . Skog 8: J . Skog l i 0 4 (BH, E, NY, S); Rio Maricao above Maricao, 20 October 1913, F. Stevens 8: W . Hess 3716 (KY); Maricao, 18 Sovember 1913, F. Stevens 8: W. Hess 4872 ( S Y ) ; one-third of way down path from El Monte de Estado to fish hatchery at Maricao, 2000 ft, February 1965, T. Talpey 30 (BH, US). LOCALITY UNKNOWN: matted in mountain brook, forest, 19 May 1935, F . Snrgent 613 (US). CULTIVATED: Cornell Univer- sity, 15 January 1964, R. Clark s n (BH). DIscussIoN.-Gesneria pauciflora resembles Ges- nel-ia humilis from Haiti and Cuba in its vegeta- tive and fruit characters, and habitat. Gesneria humilis differs in having a yellow-green to white corolla, about 1.2 cm long, a patent or reflexed limb, and greenish stems and peduncles. Gesnel-ia citrina, a closely related species in Puerto Rico, differs in having its peduncles much shorter than pedicels, leaves obovate or spathulate with rounded apices, in its decumbent habit and drier habitat. From among the many duplicate specimens of the holotype (Sintenis 327), which was at Berlin and probably destroyed, a specimen at Gottingen has been selected as a lectotype. This specimen bears flowers and fruits, and the sheet was anno- tated by Urban as Gesneria pazicipora. Section 6. Physcophyllon L. Skog, new section Plantae erectae vel decumbentes acaulescentes, caulibus raro usque ad 0.5 m altis, ex basi vel ex foliorum superiorum axillis ramificantes. Folia cuneata, lanceolata vel oblanceolata, membrana- cea, bullata, scabrosa vel pilosa, marginibus inter- dum crispis vel lobulatis. Inflorescentiae 1- co- florae, quam foliis subtentibus breviores; flores tubulosi, fere cylindracei, curvati vel subventri- cosi, aurantiaci vel rubri, staminibus inclusis. Capsulae turbinatae vel fere sphaericae, interdum calycibus praeditae scyphulas formantes, in iMar- chantiae (bryophytorum) memoriam revocantes, costis prominentibus vel obscuris. TYPE-SPECIES.-Gesneria cllneifolia (A. P. de Candolie) Fritsch. DIscussIox.-Section Physcophyllon comprises a complex of species, occurring on the islands of the Greater Antilles, which share similar characters and resemble each other to such a degree that they are distinguished with difficulty. This section resembles sections Gesneria and Chol-isanthel-a in habit, foliage and fruit characters, and may be closely related to them. Because of the presence of the red tubular corolla in all of the species, the pollinator of each of the species is probably a hummingbird. Name derived from the Greek physke (= blis- ter) + phyllon (= leaf). 27. Gesneria acaulis Linnaeus Gesneria acaulis Linneaus, Syst. Nat. ed. 10, 2:1110, 1759. Subshrubs: stems woody, erect, pendent or de- cumbent, short, usually about 20 cm, rarely to 60 cm tall, 3-5 mm in diameter at 5 cm below apex, bark smooth and gray-brown to glabrescent and greenish toward the resinous apex, lenticels ob- scure or erumpent longitudinally; rarely branched or branching from the base, internodes 0.1-2.0 cm long, leaf scars prominent, Leaves a1 ternate, spiraled, clustered at the apex of stem: petioles sulcate, 0.1-1.9 cm long, 0.1-0.3 cm wide, green or appearing reddish from red- dish multicellular trichomes, villous and some- what resinous from short glandular trichomes; blades oblanceolate to obovate or often falcate, (2.5-)3.6-23.4 (-25.0) cm long, 1.0-8.1 cm wide, membranous, base subcuneate to subcordate, usu- ally oblique, margin crenate, dentate, serrate, double-serrate or irregularly lobulate, apex acu- minate, acute or obtuse, adaxial surface bullate to NUMBER 29 Key to the Species of Section Physcophyllon 1. Petioles, floral tubes, and capsules verrucose; calyx lobes connate for ca 3 mm: Haiti ................ 1. Petioles, floral tubes, and capsules smooth or pubescent, not verrucose; calyx lobes connate for 2 mm or less. 2. Leaves narrowly oblanceolate, margins regularly undulate-lobulate; calyx lobes narrow, 1-2 mm wide a t base, apex recurved; inflorescences usually of one flower; peduncles 2 G. christii 2 . Leaves oblanceolate or obovate, margins irregularly serrate or crenate; calyx lobes usually broad, if narrow then apex not recurved; inflorescences of one to many flowers; peduncles usually longer than 2 mm. 3. Corolla tube straight, or only slightly curved at apex, cylindric, limb patent, lobes nearly equal; Cuba, Hispaniola, and Puerto Rico ........................ 35. G. reticdata 3. Corolla tube curved at or near the middle, usually ventricose, if not ventricose then leaves glabrous above, limb spreading or lobes erect or reflexed and usually unequal. 4. Subtending leaves ca 5 times as long as the inflorescences, leaves spathulate or 4. Subtending leaves either less than 4 times as long as the inflorescences, usually much less, or leaves irregularly serrate. 5 . Corolla 1.5-1.8 cm long; peduncles glabrescent; flowers many; southern Domini- 5. Corolla 1.8 cm or more long, if less than peduncles pubescent to villous; flowers one to many. 6. Leaves scabrous; persistant calyx lobes on capsule reflexed, peduncles 0.8- 6. Leaves bullate or Emooth, if scabrous then persistant calyx lobes erect or spreading, peduncles 0.1-3.1 cm long. 7 . Capsules obconic or narrowly turbinate; leaves bullate or smooth, not scabrous; peduncles 0.1-0.5 cm long, rarely to 2.6 cm; bracts 0.3-1.0 cm long; inflorescences of 1-3 flowers; Puerto Rico .............. 30. G . cuneifolia 7. Capsules broadly turbinate or globose; leaves scabrous or not: peduncles 0.2-3.1 cm long; bracts to 3.4 cm long, usually more than 1 cm long; inflorescences of 1-5 or many flowers. 8. Calyx lobes erect, 1.5-7.0 mm long; inflorescences occasionally as long as the subtending leaves; corolla scarlet or dark red, base attenuate and ventricose above; leaves scabrous; Cuba . . . . . . . . . . 32. G. libanensis 8. Calyx lobes spreading, 7.0-20.0 mm long; inflorescences shorter than the subtending leaves; corolla red, dull rose or orange, base ampliate and not ventricose above, or narrow at base and ventricose above; leaves seldom scabrous, but glabrous in var. glabrata; Jamaica. . . . . . . ........................................................................................................ 27. G . acaulis ......................................................................... ....................................................... 31. G. hybocmpa mm long or less; Haiti ............................................................................... 29. narrowly obovate, margin crenate; Cuba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34. G. purpurascens .................. 28. G . burahonensis can Republic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.2 cm long: northern Dominican Republic .................... 33. G. pedicelluris 101 flat, light to dark green, villous or trichomes stiff and appressed, trichomes occasionally suffused with calcium carbonate and whitish, or glabrous, veins immersed, abaxial surface lighter green with prominent dark green or reddish veins, villous or trichomes appressed usually more dense at the veins, resinous from glandular trichomes. Inflorescences 1-3 (-many) in pedunculate cymes about one-quarter of the length of the subtending leaves: peduncles terete, 0.1-3-1 cm long, 1-2 mm in diameter, green or red, puberulous to villous, occasionally resinous; bracts leaflike, linear, lance- olate, or elliptic, to 3.4 cm long, to 0.8 cm wide, green or red, villous to glabrescent; pedicels terete, 0.4-2.9 cm long, elongating in fruit, 1-2 mm in diameter, green or red, glabrous or puberulous to villous; floral tube obconic or turbinate, 2-4 mm long, 1.2-3.0 mm wide, green or red, villous to glabrescent, resinous, costae 5 or 10, usually prominent; calyx lobes 5, spreading, connate at base for 0.5-2.0 mm, each linear or lanceolate, 0.7-2.0 cm long, 1-4 mm wide at the base, narrow- ing gradually to apex, margin entire, outer surface green or red, glabrous to villous, occasionally villous only near the margin, trichomes many- celled, or appressed, stiff and white, glandular- S!dITHSOEIAN CONTRIBUTIONS TO BOTANY resinous, veins 3-5, nearly parallel, inside green or red, glabrous or sparingly pubescent, glandular- resinous, glands becoming more dense toward the base; corolla tube cylindric or subventricose, bent at or above the middle, little or much expanded above floral tube, 1.8-3.6 cm long, 2-4 mm in diameter, middle 4-7 mm in diameter, narrowing somewhat at the throat, 3-6 mm in diameter, outside red, dull rose, or orange, occasionally yellow at the veins, puberulous to villous, tri- chonies occasionally of 2 lengths, inside yellow or light red, glabrescent or glabrous, limb 5-lobed, nearly erect, darker red-orange, upper lobes ex- ceeding lower lobe, upper lobes broadly ovate, 1.0- 2.5 mm long, 2.0-3.0 mm wide, margins suberose, stipitate-glandular, lateral lobes and basal lobe broad, rounded, seldom more than 1 mm long and 2 mm wide; stamens 4, adnate for about 1 mm to base of corolla tube, exserted only to length of upper corolla lobes, filaments linear, curved, 1.8-3.5 cm long at anthesis, less than 1 mm in diameter, reddish, yellow, or white and nearly transparent, sparingly pubescent with a few scat- tered trichomes near the middle, anthers oblong, 1-2 mm long, 1 mm wide, pinkish, yellow, or white, glabrous, adherent in 2 pairs, staminode up to 5 mm long with undeveloped anther; ovary com- pletely inferior, apex villous, disc 5-angled, un- lobed, glabrous, style linear, curved, slightly ex- ceeding the stamens at maturity, red, yellow, or white, sparsely pubescent, stigma clavate, recurred, yellow or white, papillate. Capsule broadly turbinate to globose, 2-4 mm long, 4-6 mm wide, grayish brown or gray, villous or glabrescent, dehiscing from the apex, costae 5-10 and usually prominent; seeds numerous, fusiform, twisted ca 0.5 mm long, ca 0.25 mm wide, reddish-brown to black, nitid (Figures 19b, e). 27a. Gesneria acaulis Linnaeus var. acauZis Gesneria acaulis Linnaeus, Syst. Nat. ed. 10. 2:1110, 1759.- Linn., in Elmg., PI. Jam. Pug. 15, 1759.-Linn, Amoen. Acad. 5:400, 1760; Sp. PI. ed. 2. 23850, 1763; Syst. Nat. ed. 12. 2:409, 1767.-Murr., Syst. Veg. ed. 13, 554, 1784-Sw., Obs. Bot. 227, 1791.-Poir. in Lam., Illustr. 3:88, 1823 [as to description, not as to illustrated plant].-Urb., S7mb. 4 n t . 2:379, 1901.-Adams, F1. PI. Jamaica 680, 1972. Conradia sloanei A. P. de Candolle, Prodr. 7:526, 1839.- Griseb., F1. Brit. W. Ind. 462, 1862 [nom. illeg.]. [Type- collection: Jamaica, 0. Szcartz, sn (G-DC, holotype; B, C, LD, M, S, UPS, isotypes).] Conradia hispida Bentham, P1. Hartw. 264, 1846.-Walp., Repert. Bot. Syst. 63395, 1847. [Type-collection: Between Kingston and Spanish Town, Jamaica, C. Hartweg 1551 (K, holotype; BM, K, LD, isotypes).] Chorisanthera hispida (Bentham) Qrsted, Cent. Gem. B, 1838. Conradia libanensis sensu Grisebach, F1. Brit. \V. Ind. 4@, 1862 [excl. syn.]. Pentarhaphia h isp ida (Bentham) Hanstein, Linnaea 34:293, 1865. Pentarhaphia sloanei (?4. P. de Candolle) Hanstein, Linnaea 343295, 1865.-Hitchcock, Rep. Missouri Bot. Gard. 4:114, 1893 (?Pentaraphia?). Gesneria hispidn (Bentham) 0. Kuntze, Rev. Gen. 2:473, 1891. Pentarhaphia acaitIis (Linnaeus) Bentham & Hooker f . ex Fawcett, Prov. List, Jam. 28, 1893.-Druce, Rep. Bot. Exch. C1. Brit. Isles 3:422, 1913 [?1914?]. [?Pentarraphia?]. Pentarhaphia libanensis (Grisebach) Bentham & Hooker f. ex Fawcett, Prov. List, Jam. 28, 1893 [nom. illeg., non Hanstein (1865)]. Gesneria acaiilis Linnaeus var. grandifolia Urban, Symb. Ant. 2:380, 1901. [Type-collection: Jamaica, C. Bertero 2183 (TO, lectotype).] Leaves pubescent above, margin lobate, often double-serrate, apex acute to obtuse, stomata scattered, seldom in Lgroups. Calyx lobes bearing many-celled or appressed trichomes on the exterior at least at the margin, inside glandular; corolla tube cylindric, inflated at base, ventricose at middle, narrowed and bent slightly down above middle, slightly flared at apex; pollen grains isopolar, prolate, size small (18.5- 22.5 pm long at the polar axis, 10.2-12.5 pm wide at the equatorial axis), amb nearly circular, tri- colpate, apocolpia truncate or slightly rounded, sexine heterobrochate, lumina 0.4-1.5 pm across (Figure I7d). TYPE.-H. Sloane, Voy. Jam. Hist. 159, pl. 102: fig. 1, 1725 (Figure 60). CHROMOSOME NUMBER.--^ = 14 (Lee, 1966), and fide Davidse & Conroy 3270 (MO). var. acaulis is endemic to Jamaica (Figure 61), where i t can be found in all parishes except Man- Chester. Its habitat is damp rocky soil or banks from ca 25-700 m elevation. This variety flowers throughout the year. DISTRIBUTION AND Eco~oc~.-Gesnerkz aCUUl i5 SPECIMENS ExA?+IIxED.-JAMAICA. P.ARISH OF HANOVER: 5 mi TV of Lucea, near sea level, 28 February 1968, G. Proctor 28559 (F, IJ, MO). PARISH OF WESTMORELAND: One-day Cave, NUMBER 29 103 Key to the Varieties of Gesneria acaulis 1. Leaves often double-serrate, pubescent above; calyx lobes bearing many-celled or appressed 27a. var. acuulis 1. Leaves never double-serrate, but crenulate to dentate or single serrate, glabrous above; exterior of calyx lobes glabrous: corolla bent at middle, scarcely wider than the base at any trichomes a t least at the margin dorsally; corolla bent above the ventricose middle . . . . . . . . . . . . ............................................................................................................................................. point ..................................................................................... 27b. var. glabrata, new variety ca 1 mi W of Rat Trap, 1000-1100 ft, 9 April 1961, G. Proc- tor 22160 (BM). PARISH OF ST. ELIZABETH: Cook?s Bottom, N of Ipswich, 40C-450 m, 31 March 1920, W. Maxon Fi E . K i l l i p I460 (A, F, GH, NY, US). PARISH OF TRELAWNY: 1 mi SW of Bunkers Hill, ca 400 ft, 12 January 1964, G. Proctor 24493 (BM, IJ); Windsor House estate, 400 ft, 11 March 1956, W . Stearn 469 (A, BM); path to Windsor Cave, 450 ft, 23 August 1955, D. Powell 140 (IJ); SW of Windsor caves, ca 500-750 ft, 13 August 1963, M . Crosby & W. Anderson 1171 (F, MO, NY, US, USF); Burnt Hill, 500 m, 29 April 1952, F . Barkley 223236 (IJ, US); 0.5 mi N of Spring Garden, ca 1700 ft, 4 November 1968, G. Proctor 29329 (IJ); Barbecue Bottom, 6 mi N of Albert Town, Cockpit Country, 29 July 1962, F . Fosberg 42920 (NY, Us). PARISH OF CLARENmN: Peckham Woods, 2000-2250 ft, 10 July 1960, C. Adams 7545 (UCWI). PARISH OF ST. ANN: near Brown?s Town, 18 January 1938, F . Hunnewell 15356 (GH); Liberty Hill, 27-30 March 1908, N . Britton 2500 (NY); 2.2 mi W by road of Albion, 2300-2400 ft, 9 August 1965, H . Hespenheide 923 (GH, MO, KY, US, Z); Dry Harbour Mountains, 2 mi W of Albion, 18 January 1956, W . Stearn 155 (BM, 2 sheets); Roar- ing River Falls, 7 March 1936, F. Hunnewell & L . Griscom 14386 (IJ); near Lydford PO, 1500 ft, 20-31 December 1953, R . Howard & G. Proctor 13444 (A, IJ); Union Hill, near Moneague, 13 July 1950, R . Howard 12017 (A, US); Union Hill and vicinity, northern slopes of Mount Diablo, 400-750 m, 24 July 1926, W. Maxon I0424 (GH, US); Moneague, 27 February 1927, E . Wall sn (S); Moneague, December 1849, R . Alexander sn (GOET); Union Hill, near Moneague, 650 m, 6-7 April 1908, N. Britton Fi A . Hollick 2804 (NY); Roar- ing River Falls, 27-30 March 1908, N. Britton 2487 (NY); Dunn River Falls, 11 March 1936, F . Hunnewell & L. Gris- corn sn (GH); Hollymount Road, 2300 ft, 27 August 1963, C. Adams 12644 (BM, UCWI); Mt. Diablo, 2 April 1903, W . Harris 8485 (UCWI); Mt. Diablo, 8 March 1936, F . Hunnewell & L. Griscom 14385 (GH); eastern slopes of Mt. Diablo, 7 March 1916, E . Killip 448 (BM, MO); Mt. Diablo, January 1850, R . Prior 560 (K); Mt. Diablo, 1800 ft, 20 March 1970, M . d u Quesnay 257 (UCWI); Mt. Diablo, 2000- 3000 ft, March 1952, R . Robbins 1513 (UCWI); along road to Hollymount above Mount Diablo Afforestation area, 13 August 1970, L. Skog 1634 (BH, US); Mt. Diablo, road to Hollymount, ca 1.7 mi SW of Schwallenburgh, 11 July 1954, G. Webster Fi K . Wilson 5017 (A); on way to Hollymount, 2000-2500 ft, 18 February 1958, T. Yuncker 18215 (BM, F, s); Schwallenburgh, Mt. Diablo, ca 2000 ft, 30 November 1964, D. Powell 1642 (IJ); between Mount Rosser and Schwallenburgh, on east side of Mount Diablo, 1700 ft, 12 FIGURE 6O.-Type of Gesneria acaulis Linnaeus: Sloane, Voy . Jam. Hist., pl. 102: fig. 1, 1725. ShlITHSOSIAS C O S T R I B U T I O S S T O BOTAKY April 1956, W . Stearn 712 (Bhl): Moneague to hfount Diablo, 5 April 1908, A?. Britton 2720 (F, NY); cliffs in Fern Gully, 19 March 1897, J . Churchi l l sn (GH); Fern Gully, February 1916, H . Ridley sn (K): Fern Gully, 500-1000 ft, July 1951, R. Robbins 1517 (UCFVI); along Hwy -43 in Fern Gully about 3 mi from Ocho Rios, 13 August 1970, L . Skog 1633 (BH, US); near Fern Gully, above Ocho Rios, 500-1000 f t , 23 December 1957, T . Yuncker I7781 (BM, F, S, CCTVI): nr. Burts Run, 1800 ft, 13 Xovember 1965, C . A d a m 12745 (CCT\?I); Somerton district, ca 1700 ft, 28 March 1956, G. Proctor 8; JV. Stearn 11923 (IJ), near hlosely Hall Cave, 1 mi T V of Blackstonedge PO, ca 2000 ft, 15 July 1952, G. Proctor 6926 (IJ, SY). PARISH OF ST. CATHERINE: Forest Reserve area east of Crofts hlountain, 1650-1850 ft, 6 September 1962, G. Proctor 22762 (BM, IJ); Holly Mt., 2 August 1931, L. Bailey 705 (BH): Hollymount, 19, 20 September 1906, A?. Britton 733 (NY); Hollymount near Ewarton, 2500 ft, 10 August 1896, U?. Harris 6453 (Bhl, C, UCTVI), 14 February 1905, I??. Harris 8886 (Bhl, F, K, S Y , UCTVI), 31 August 1905, TV. Hnr- ris 900? (BM, F 2 sheets, K, S Y , UCTVI): Hollymount and vicinity, 750-850 m , 24 July 1926, W . A f a x o n 10449 (S, US); Holly Mount, 2600 ft, February 1916, J . Perkins 487a (G); 1 mi due 1%? of Riverhead, ca 1500 ft, 16 October 1960, G. Proctor 2Iff3 (BM, IJ, US); Linstead, 400 ft, 1 Sovember 1953, G. Proctor 8148 (IJ, USF); Linstead, December 1959, I.. Wynter 5669 (UCTVI); Bog T\?alk, 4-5 May 1910, J . Craw- ford 832 (SY, PH); 2.1 mi SE of Bog Tt?alk along main hwy to Spanish Town, 80 m, 26 Sovember 1971, G. Davidse .k E . Conroy 3270 (MO); Bog TValk, 2 February 1901, W. F a u w t t 8127 (BM, NY); Bog TValk, 17 December 1890, A . Hitchcock sn (F, MO); Bog TValk and vicinity, 27 July 1926, TY. Maxon I0485 (S, US); Bog Tt?alk, 10 March 1885, A?. Kidder $71 (GH, 2 sheets); Bog TValk, February 1916, H . Ridley sn (K); Bog TValk, 17 December 1890, J . Rothrock 87 (F); Flat Bridge to Bog TValk, 200 ft, 13 March 1960, C. A d a m 6476 (MO, UCTVI); Rio Cobre, river bank south of Kent Village, ca 250 ft, 30 June 1963, JI. Crosby, H . Hespenheide .k W . A n - derson 410 (F, S Y , CSF); cliffs near Rabys Corner, ca 2 m i SE of Bog T\?alk, ca 500 ft, 10 June 1959, G. Webster, J . Ellis .k K. Miller 8087 (BM, G, S); Natural Bridge, Riversdale, 400-500 ft, 3 April 1961, G. Proctor 22145 (BM, IJ); between Kingston and Spanish Town, C. Hartweg 1551 (K, holotype of Conradin hispida Bentham: BM, K, LD, isotypes); St. Thomas in the Vale, June 1843, TY. Ptirdie sn (K); on the road to Sixteen-Mile-TValk, H . Sloane sn [Horti Sicci 3:27] (BM). PARISH OF ST. MARY: Dressikie, 600-700 ft, 3 xovem- her 1968, G. Proctor 29306 (IJ); 1 mi S of Boscobel PO, 300 ft, 2 May 1952, G. Proctor 6614 (IJ, US). PARISH OF ST. AXDREW: Iron Face, Chester Vale, 3000 ft, 2 December 1907, IV. Harris 10026 (A, BM, C, F, K, S Y , P, UCTVI, US, 2): vicinity of Cinchona, Old England, 2-10 September 1906, D. j\Jarble 242 (NY): Port Royal Mountains, November 184.5, G. MacA?ab sn (P), no date, W. Lane 443 (K): in montibus (copia pluviale minore) prope Kingston, 1897, C. Hanson $12 (C, L). PARISH OF PORTLAND: near Chester \?ale, 9 December 1891, 1.V. Fau?cett sn (UC\\?I); Haycock Hill, 18 March 1962, A . Skelding in C . A d a m 10857 (UCTVI); along road between Section and Silver Hill Gap, ca 3500 ft, 3 December 1963, G. Proctor 24272 (BM, IJ); John Crow Peak, n o date, J . Hart I219 (UCTVI); vicinity of Thomson?s Gap, ca 1000 m , 5-6 March 1920, U?. h faX0?7 & E. Killip 785 (US): Blue Mts. between Vinegar Hill and Thomson?s Gap, 30 March 1916, J . Perkins 1025 (GH); S e w Eden, January 1847, J . m?olle sn (USF): Swift River Gorge a t Eden, y2 mi E of Paradise, 100 ft, I9 March 1956, 1%?. Stearn 552 (BM); Swift River Bridge near Paradise, ca 80 f t , 25 Sovember 1959, C. Adonis 5570 (UCTVI): St. Margaret?s Bay, Sovember 1900, C. l l f i l lspa~igl~ 19-77 (F); near mouth o f Rio Grande, ca 25 ft, I9 March 19i6, G . Proctor 8; TV. Steclrn 11859 (IJ): mouth of the Rio Grantle, 20-30 ft, 18 March 1956, TY. Stearn 546 (BM. 2 sheets); St. Margaret?s Bay, 5 mi T?V of Port .4ntonio, mouth of the Rio Grande, 18 February 1906, A . Wight 130 (F, S Y ) : coast road near Port Antonio, 9 May 1904, TV. , l fnxon 2105 ( S Y , US): near Moorestown, 25 September 1906, D . Afnrble 856 (SY); Ginger House to Comfort Castle, 400 ft, 10 August 1962, C. Adonis 11471 (CCTt?I); Corn Puss Gap ca 2000-3000 ft, 14 August 1954, K. Wilson 8; W. Mtirray 59f (A , IJ); S of Ecclesdown, 1500-2000 ft, 3 November 1964, T . T o l p y 15 (RH); l..i-2.5 mi STV of Ecclesdown, 1500-2500 ft, 24 January 1956, R. Howard, G. Proctor 8; W. Stearn I4797 (A, BM, IJ); Cuna-Cuna Pass, 24 July 1897, A . Fred- h l ~ n 3172 (SY): Cuna-Cuna Pass, 15 September 1908, W . Hnrj-i.i 8s S, Britton 10559 (F, K, S Y , UCTVI); John Crow Mt., March 1931, X. Robbins 1516 (UCTt?I); ST\? slope of Joe Hill, 1600 ft, 13 February 1956, W . Stearn 246 (A, BH, BM, K). PARISH OF ST. THoxr.ks: Rowlandsfield district, SE slope of John CroTv hits., 18 March 1952, G. Proctor 6428 (IJ, US): above Roxlandsfield, 1500 ft, 16 March 1956, W . Stearn 510 (BH, AM). LOCALITY UXKSO~VS: 1850, R. A l e x - ander s~7 (K): June 1821, C. Bertero 2183 (TO, lectotype of Gesneria acaulis var. grandifolin Urban); no date, P. Browne s n (UPS); no date, H . Distin stz (K); no date, G o u w c o k - tion s n (E): borealis in montibns humidus copia pluvialis, 1897, C . Hansoii sn (C); Dollwood, October 1893, W . Harris s n (BXI, CCTVI); no date, J . Hart srz (US); no date, IY. Lane 84 (K), March 1939, J . Roig If224 (SV); no date, J . Mac- F n d y e n sn (K): 18,58, TV. dfarclz 1197 (K); no date: 0. Swartz sn (G-DC, lectotype of Conradia sloanei A . P. de Candolle: B, C, LD 2 sheets, M, S 2 sheets, UPS, isotypes); n o date, .I. W i l e s s n (G). CUI.TI~ATED: Cornell University, G-877, 2 August 1966, I-. Leva 39 (BH), 5 October 1969, I>. Skog 1410 (BH), 3 November 1970, L . Sko,q 1732 (BH), 27 January 1967, Af. Stone 216 (BH); Cornell University, G-1003, 18 October 1966, M . Stone 23f (BH). DiscvssIozr.--Jn the protolog for Gesneria acaulis, Linnaeus cited Sloane?s plate of Rapztnculo a f in i s anomala vasculifera, . . . (?Sloan, jam. t. 102, f . 1?). This must be considered the type for the name. A specimen in the Sloane herbarium (Horti Sicci 3:27) at the British Museum (Natural History) is a match for the drawing and may be considered the typotype in the sense of Dandy (Stearn, 1957). Apparent natural bigeneric hybrids between Gesneria acaitlis and Rhytidophyllum tomentosum NUMBER 29 105 (Linnaeus) Martius have been collected in the parishes of St. Ann and Manchester. Specimens of C. Adams 12643 and G. Proctor 29384 were the basis for the separation of Gesneria sp. ?A? by Adams (1972). Characters appear intermediate between the two species, but no viable seed has been found. Hybrid plants may be expected wher- ever the species are found growing in the same locality. Plants of an apparent hybrid between Gesneria acaulis var. glabrata and Rhytido- phyllum tomentosum were introduced into culti- vation in 1970 and grown at Cornell University under the accession number G-1371 (Figure lj). The arrangement of the stomata of the abaxial leaf epidermis was discussed and illustrated by Wiehler (1970). He pointed out that the two clones of Gesneria acaulis in cultivation at Cornell University differed markedly in stomatal arrange- ment. The epidermis of a typical specimen of the variety described here as var. acaulis (from the Parish of Portland and grown under the accession number G-877) had stomata scattered more or less generally over the abaxial surface. A typical speci- men of var. glabrata (G-876, from the Parish of Manchester), described below as new, has stomata in apparent clusters (termed ?islands? by Wiehler). These clusters appear to correspond to the sum- mits of the stomatal domes. It seems likely that if the epidermis did not have the domes, the stomata would not appear to be clustered. Weihler prob- ably examined only the two specimens in culti- vation. My examination of the two cultivated clones found the cells arranged as he described; however, there appears to be considerable varia- tion in samples collected from the area between the typical forms, particularly in samples from the Cockpit Country of Jamaica. Some examples bear one to few stomata in an apparent cluster perhaps at the apex of a stomatal dome. 27b. Gesneria acaulk var. glabrata L, Skog, new variety Folia supra glabra vel prope basim raro sparsim pubescentia, ad margines crenulata vel dentata vel singulatim serrata, non lobata, ad apices acuta vel acuminata, cellulis epidermalibus plerumque dis- cretis. Calycis lobi utroque dense glandulosi, venis 3 prominentibus; corollarum tubi cylindracei, non ventricosi, ad medium deorsum flexi (Figure 16g). TYPE-CoLLECTIoN.-Marshall?s Pen, Jamaica, L. Skog 1642 (US, holotype, Figure 62a; BH, isotype). CHROMOSOME NUMBER.-TZ = 14 (fide Skog 1731). DISTRIBUTION AND EcoLocz.-Gesneria acaulis var. glabrata is known only from the parish of Manchester in Jamaica (Figure 61) where it grows on shaded limestone banks at 650-800 m elevation. This variety flowers throughout the year. SPECIMENS ExAMINED.-JAMAICA. PARISH OF MANCHESTER: Bellefield to Banana Ground, 2450 f t , 11 December 1960, C. Adams 8451 (UCWI): Mandeville and vicinity, 29 August FIGURE 61.-Distribution of Gesneria section Physcophyllon in Jamaica. (G. acaulis var. acaulis = open circles: G . acaulis var. glabrata = solid circles.) I06 SMITHSOSIAN CONTRIBUTIONS TO BOTANY 1907, N. Britton 1000 (NY); cliffs, Somerset, 21-23 Septem- ber 1908, N . Britton 3732 (NY); Mandeville and vicinity, 15-26 February 1910, S. Brown 68 (NY, PH), 239 (NY, PH, US); Marshall?s Pen, Mandeville, 2200 ft, 29 April 1896, W . Harris 6322 (BM, UCWI, US); Walderston, 2200 ft, 14 Octo- ber 1901, W . Harris 8415 (BM, UCWI); Marshall?s Pen, no date, A . Katrenberger & R. Katzenberger 113 ( R D J ) ; vicinity of Marshall?s Pen Estate, near Mandeville, 2300 ft, 24 April 1961, K. Kramer 1691 (U, WAG 2 sheets); Cockpit Country, bank near Derry, 13-18 September 1906, D . Marble 709 (F, NY, US); Marshall?s Pen, 2.25 mi due SW of Mandeville, 2100-2300 ft, 16 August 1965, G. Proctor 26617(IJ); Mar- shall?s Pen, 6 January 1959, E . Robertson 5374 ( K , UCWI); Marshall?s Pen, 2.25 mi NW of Mandeville, 2100-2300 f t , 15 August 1970, L. Skog 1642 (US, holotype of Gesneria acaulis var. glabrata L. Skog; BH, isotype); Somerset, 2300 ft, 23 February 1956, W . Stearn 350 (BH, BM, K); S of Green Hill, 5 November 1964, T. Talpey 18 (BH); halfway between Coleyville and Molton, 2000-2250 ft, 20 June 1939, G . W e b - ster, ]. Ellis & K. Miller 8421 (BM, G, s); Bethany, 1849, H . WdlschlGgeE 957 (M). LOCALITY UNKNOWN: 1855, T. Hogg sn (NY); no date, J . MacFadyen sn (K); no date, W . Wright sn (BM); no date, H . Wullschlugel sn (GOET, W). CULTI- VATED: Cornell University, G-876, 5 October 1969, L. Skog I425 (BH), 3 Eovember 1970, L. Skog 1731 (BH). DIscussIoN.-This variety was brought into culti- vation by T. Talpey in 1964 and was grown at Cornell University as G-876. 28. Gesneria barahonensis Urban Gesneria barahonensis Urban, Symb. Ant. 7:379, 1912. Subshrubs: stems unbranched, woody, to 15 cm tall, glabrescent below, villous and resinous toward apex, internodes short, to 5 mm long. Leaves alternate, clustered near stem apex: petioles sulcate, 3-6 mm long, 1-2 mm wide, green or reddish, sparsely pilose and glandular; blades oblanceolate to obovate, 5.6-15.4 cm long, 2.4-5.1 cm wide, membranous, base acute or rounded, margin serrate to crenate, apex acute, adaxial sur- face slightly rough, green, pilose with appressed white trichomes, abaxial surface lighter green, villous and glandular along the prominent veins. Inflorescences condensed cymes, nearly umbel- late, flowers many, the whole about one-half the length of the subtending leaf: peduncles terete, 0.7-2.3 cm long, ca 1 mm in diameter, glabrescent; bracts 2, linear, 5-7 mm long, ca 1 mm wide, reddish, pilose; pedicels terete, to 1.8 cm long, ca 1 mm in diameter, reddish; floral tube obconic or turbinate, ca 2 mm long and wide, reddish to green, glandular and pilose, trichomes more dense just above junction with pedicel, veins not promi- nent; calyx lobes 5, erect, connate 1-2 mm at base, narrowly triangular, 5-9 mm long, 1-2 mm wide at base, both sides green or somewhat reddish, glandular-glabrescent at the base externally, mar- gin ciliate, 3-veined; corolla tube cylindric, but slightly broader below the middle, then narrowed toward throat, with limb slightly flaring, tube 1.5- 1.8 cm long, ca 2 mm broad at base, 4-6 mm wide at broadest point, ca 2.5 mm wide at throat, dark red outside, pilose, trichomes more dense at mouth, inside lighter red, glabrous, mouth slightly oblique, lobes crescent-shaped about 1 mm long, margin entire to erose, glandular-ciliate: stamens 4, with a reduced staminode, adnate to base of corolla tube, exserted only to corolla limb, fila- ments slightly curved, glabrous, reddish, anthers oblong, ca 1 mm long, ca 0.5 mm wide, slightly coherent; ovary inferior, disc annular, lobed, style curved. equaling the stamens in length, red, stigma clavate. Capsule obovoid, 4-6 mm long, 3-4 mm in diameter, gray-brown, glabrescent, costae 10; seeds fusiform, twisted, ca 0.5 mm long, dark brown or black. TYPE-COLLECTION.-?An Felsen der Tha?ler von Pae-Mingo nach Bahoruco,? Dominican Republic, M . Fuertes 1049 (A, lectotype, Figure 62b). DISTRIBUTION AND EcoLocY.-Gesnerin baraho- nensis is apparently endemic to the Barahona Peninsula of Hispaniola (Figure 54), where it grows on rocky cliffs from 100-1200 m elevation. This species was collected in flower in September. SPECIMENS EXAMIh?ED.-HISP,4NIOL.4, DOhlINICAh? R E P U B - LIC: Barahona Province: an Felsen der Tha?ler Yon Pae Mingo nach Bahoruco, 100-1200 m, June-September 1911, M . Fuer- tes 1049 (.4, lectotype of Gesneria barahonensis Urban); in saxosis convallium inter Pae-Mingo et Bahoruco, 100-1200 m, 17 September 1912, ill. Fuertes 1399b (A, E, NY, P). DIscvssIoN.-This species is known only from two collections, both made by Fuertes in probably the same locality. The type, Fziertes 1049, consists of a single sheet annotated by Urban in the Arnold Arboretum, which is designated the lectotype until an authentic holotype is found. This sheet, unfor- tunately, is a mixed collection: two specimens are truly Gesneria barahonensis, the third is a speci- men of G. reticzilata (Grisebach) Urban NUMBER 29 107 FIGURE 62.-Type specimens: a, holotype of Gesneria acaulis var. glabrata L. Skog, new variety, Skog 1642 (US); b, lectotype of Gesneria barahonensis Urban, Fuertes 1049 (A). (Lectotype includes specimen on right and lower plant.) lacking flowers and superficially resembling G . barah onensis. Gesneria barahonensis is a rare 'species closely related to other members of this section and most similar to G . acaulis Linnaeus of Jamaica and G. christii Urban of Haiti. This species differs from both in having a much shorter corolla tube, narrower sepals than G . acaulis and many more flowers per inflorescence than G . christii. 29. Gesneria christii Urban Gesneria christii Urban, Symb. Ant. 6:42, 1909. Gesneria acauZis sensu Poiret, in Lam., Illustr. 3:88, pi. 536: fig. 1 , 1823, non Linnaeus (1759). Subshrubs: stems slow-growing, woody, erect or pendent, up to 12 cm tall, thick, to 8 mm in diam- eter at 5 cm below apex, bark smooth or rugose, brown and glabrescent below, above green, sparsely pilose, apex resinous; branching from base and above, internodes short, usually less than 6 mm long, leaf scars obvious. Leaves alternate, spiral, closely spaced: petioles sulcate, 5-9 mm long, 1-2 mm wide, green or reddish, pilose or villous; blades narrowly obovate or narrowly oblanceolate, occasionally falcate, 6.3-26.9 cm long, 0.7-4.9 cm wide, membranous, base narrowly cuneate or acute, margin regularly undulate-lobulate, lobules serrate or dentate, apex acute, adaxial surface bullate to flat, light to dark green, with sparse appressed whitish trichomes from enlarged bases, lateral veins diverging at an 108 S.\IITHSOSIAh' COSTRIBUTIOSS T O BOT.4SY angle usually less than 45O from the midvein, abaxial surface much lighter green with prominent reddish veins, trichomes very sparse, appressed, veins pilose with numerous short glands among the eglandular appressed trichomes. Inflorescences of usually one maturing flower, rarely 2 or 3 in cymes, much shorter than the sub- tending leaves: peduncles very short, at most 2 mm long, ca 1 mm in diameter, green or red, glandular- resinous; bracts 2, linear, ca 1 cm long, green, sparsely pilose; pedicels terete, 1.0-4.2 cm long, elongating in fruit, ca 1 mm in diameter, green or red, pilose and glandular; floral tube ovoid, 2-3 mm long, 2-3 mm in diameter, green or red, villous; calyx lobes 5, erect, connate at base for 1-2 mm, linear or lanceolate, 0.6-1.6 cm long, 1-2 mm wide above base, narrowing rapidly to middle, then gradually to apex, apex recurved, curling on drying, 3-nerved, margin entire, outer surface FIGURE 63.-Lcctotype of Gesneria christii Urban. Christ 1888 (IJ). green with reddish veins, pilose, glandular, inside green, glabrous or very sparsely pilose, but glandu- lar; corolla tube ventricose or inflated at the mid- dle, narrowed at both ends, bent upward above the base, and slightly downward (Figure 16e) above the middle, 2.8-3.7 cm long, 2-3 mm in diameter at base, 5-9 mm wide at middle, 3-5 mm in diam- eter at throat, outside J ellow-orange to red, darker at the veins, sparsely pilose, glandular, inside yellow or reddish, glabrous, mouth oblique, limb 5-lobed, lobes erect to porrect, orbiculate, ca 3 mm long, 4 mm wide, darker red than tube, lighter spots at the notches, margins stipitate glandular, upper lobes erose, others subentire; stamens 4, with a reduced staminode, filaments adnate to base of corolla for only 0.5 mm, linear, slightly curved, as long as corolla tube and not exserted, white, gla- brous, anthers oblong, ca 2 mm long, ca 1.5 mm wide, yellow, coherent in pairs or free, pollen grains isopolar, prolate, size small (22.5-24.5 pm long at the polar axis, 11.0-14.0 pm wide at the equatorial axis), tricolpate, colpi 19.4 pm long, apocolpia blunt, sexine finely reticulate, lumina ca 0.5 pm wide; ovary inferior, apex green, pilose, disc thin, 5-lobed and angled, yellowish, style linear curved, as long as the corolla tube, white, stigma bilobed, recurved. Capsule ovoid or globose, 4-7 mm long, 5-7 mm wide, gray to brown, glabrescent, dehiscing from the apex, costae 10, prominent or obscure; seeds fusiform, twisted, 0.5-1 mm long, blackish (Fig- ures 19c, 20a). TYPE-CoLLECTIoN.-Furcy, Haiti, E . Christ 1888 (IJ, lectotype, Figure 63). CHROMOSOME SLXBER.-TI = 14 (Lee, 1966a). DISTRIBUTION AND EcoLocY.-Gesneria christii is known only from southern Haiti, where it grows on shaded limestone cliffs on river banks at ca 60-1500 m elevation (Figure 54). This species has been collected in the wild with flowers during January, February, July, and August. Under greenhouse conditions it flowers most of the year. SFECIVEhS Ex.\\rt\ED.-HISPASIOLA. HUT[: Dkpartcmcnt du Sud: Massif dc la Hotte, group Morne Rochclois, Mira- goanc, at Lebrun, ca 500 m, 25 July 1926, E . Ekman H6517 (C, EHH, IJ, S, US). Departcmcnt de l'Ouest: Massif de la Selle, Cr&te-i-Piquants, Port-au-Prince, sccc. Laval, ca GOO m, 28 Januar) 1926, E . E k m a n H5475 (EHH, S, US); Massif de la Sellc, Port-au-Prince, Ribihrc Froide, ca 150 m, 29 Janu- NUMBER 29 ary 1926, E. Ekman H.5480 (A, F, IJ, S), 2 August 1926, E. Ekman H6629 (EHH, G, GH, IJ, K, NY, S, Us), 21 July 1970, L. Skog, T. Talpey & D. Pfister 1608 (BH, US), 19 Feb- ruary 1966, T. Talpey 43 (BH); Furcy, 1500 m, January 1899, E . Christ 1888 (IJ, lectotype of Gesneria christii Urban); Massif de la Selle, Port-au-Prince, Morne de l?H8pita1, 400 m, 20 May 1927, E. Ekman H8192 ( S ) . LOCALITY UNKSOWN: no date, Lamarck Herb. hro. 198 (P-LA). CULTIVATED: Cor- nell University, G-1008, 4 November 1966, M. Stone 23.5 P H ) . DIscussIoN.-The holotype specimen of Christ 1888 was probably at Berlin and destroyed. The only known extant sample of the type is a leaf fragment now deposited in the Institute of Ja- maica, but originally part of the Buch herbarium in Haiti. Until other examples are found, this fragment must be the Iectotype. A specimen in the Lamarck herbarium (no. 198) in Paris, possibly collected by J. Martin, who was a gardener at the botanical garden in Port-au-Prince, is presumably the basis for Gesneria acaulis sensu Poiret. Gesneria christii Urban is apparently most closely related to G. acaulis Linnaeus of Jamaica. The specimens of G. acaulis from the eastern end of Jamaica bear a striking resemblance to G. christii in the lobulate leaves, but the leaves of G. acaulis are not regularly undulate-lobulate and are usually much wider toward the leaf apex, the peduncles are much longer, and are equal to or longer than the pedicels, and the base of the corolla tube is expanded above the apex of the floral tube. The peduncles of G. christii are very short and may not be apparent in a cursory exam- ination. The abbreviated peduncle may be the reason Urban described this species as one-flowered. On specimens which have one-flowered inflores- cences, the secondary flowers may be reduced to lateral bulges on the peduncle. 30. Gesneriu cuneifoliu (A. P. de Candolle) Fritsch Gesneria cuneijolia (A. P . de Candolle) Fritsch in Engler & Conradia cuneifolia A. P. de Candolle, Prodr. 7:526, 1839. Pentarhaphia cuneijolia (A. P. de Candolle) Hanstein, Lin- naea 34:294, 1865. Gesneria portoricensis A. P. de Candolle ex Bello, Anales SOC. Esp. Hist. R?at. 10:288, 1881, nom. nud. Conradia reticulata sensu Stahl, Estudios F1. Puerto Rico, ed. 1, 6:259, 1888; ed. 2, 3:327, 1937 [1936], non Grisebach (1866). Prantl, Nat. Pflanzenfam. 4 (3b):184, 7 August 1894. Gesneria cuneifolia Sesse & M o h o , F1. Mex. ed. 2, 144, 1894, nom. il1eg.-Urb., Symb. Ant. 4:571, 1911; 8:647, 1921 [pro partel.-N. Britt. & P. Wils., Sci. Surv. P. R. & V. I. 6:205, 1925. Subshrubs: stems herbaceous or occasionally woody, erect or decumbent, short, seldom to 15 cm tall, 3-4 mm in diameter at 5 cm below apex, apex green or red, pilose and somewhat resinous, soon becoming glabrescent; branches from the base, in- ternodes usually very short, but up to 1 cm long in rapidly elongating individuals, leaf scars promi- nent on mature stems. Leaves alternate, spiraled, usually clustered at apex of stem or branch: petioles sulcate, 0.3-1.2 cm long, ca 2 mm thick, green or reddish, pilose to glabrescent; blades oblanceolate or obovate, 2.1- 14.1 cm long, 1.3-4.2 cm wide, membranous, base acute, cuneate or subcordate, margin shallowly or grossly crenate or dentate, often ciliate, apex acute to obtuse, adaxial surface somewhat bullate, light or dark green, glabrous or rarely sparingly pilose at base or on midrib toward the base, veins lighter green, abaxial surface light green, glabrous or pilose along veins toward the base, veins promi- nent, epidermis with stomata grouped into clusters of 2 to 4. Inflorescences cymose, pendent, 1- to 3-flowered, usually shorter, but occasionally as long as the subtending leaves: peduncles short, 0.1-0.5 cm, rarely to 2.6 cm long, pubescent; bracts linear or lanceolate, green 0.3-1.0 cm long, 1-2 mm wide, pubescent; pedicels 1.4-6.5 cm long, green or reddish, pilose; floral tube obconic, longer than broad at anthesis, 2 4 mm long, 2-3 mm in diam- eter, green to dark red, costate, pilose; calyx lobes erect, narrowly triangular, connate 1-2 mm at base, 2-6 mm long, 1.0-2.5 mm wide at base, out- side green to dark red, glabrous or rarely sparsely pubescent, glandular, margin entire, ciliate, 3-5 nerved, inside glabrous, but glandular; corolla tubular (Figure 16h), obviously ventricose at middle, base gibbous on upper side, 1.6-2.6 cm long, 2-3 mm wide at base, expanding to 4-6 mm wide at middle, decreasing to 2-3 mm wide at throat before expanding at limb, outside light to dark red, with darker veins or yellowish on lower side, puberulent, inside yellow or light red, glabrous, limb of 5 lobes erect to porrect, 6-9 mm wide, red to yellow, sparsely glandular, upper lobes broadly orbiculate, 1.5-2.0 mm long, ca 3 mm 110 SMITHSOSIAN CONTRIBUTIOSS T O BOTAKY wide, serrate, lateral and basal lobes broadly ovate, ca 2 mm long, ca 3.5 mm wide, entire to suberose; stamens 4, not exserted beyond corolla throat, basally adnate to corolla for less than 1 mm, fila- ments linear, curved, ca 1.5-2.0 cm long, less than 1 mm in diameter, yellow to pink, glabrous, anthers oblong, ca 2 mm long, ca 1.5 mm wide, yellow-white, connate in 2 pairs, staminode ca 7 mm long; ovary completely inferior, sparsely pilose at apex, disc 5-angled, ca 1.5 mm wide, style curved, 1.6-1.9 cm long, white, glabrous, stigma clavate, papillate. Capsule upright, obconic or narrowly turbinate, persistent calyx lobes spreading, 4-7 mm long, 3-5 mm wide, gray, glabrous, only apex dehiscing, costae 10, prominent; seeds elliptic, striated, ca 0.75 FIGURE 64.-Type of Gesneria cuneifolia (A. P. de Candolle) Fritsch, A. de Candolle, Calq . des Dess. F l . Mex . , plate 723, 1874. mm long, ca 0.25 mm wide, tawny (Figures 19k, 20d) . TYPE.-A. de Candolle, Calq. des Dess. F1. Mex. pl. 723, 1874 (Figure 64). CHROMOSOME NUMBER.--IE = 28 (Lee, 1964; Tal- pey, 1966). LOCAL NA;MEs.-?Yerba parrera? (fide Bello), ?Flor de cueva? (fide Sintenis ex Urban). DISTRIBUTION AKD EcoLocv.-Gesneria cuneifolia grows in the central and western part of Puerto Rico (Figure 65) on shaded, damp limestone banks and cliffs at elevations of ca 250-400 m. Plants have been collected in flower during every month of the year, both from the wild and in cultivation. SPEcihiEh?s ExAMIxED.-PUERTO RICO: vicinity of Maya- guez, 4-10 March 1906, E . Britton & D. Marble 662 (NY); Finca Alvarez, Quebradillas, January 191 3, Bro. Hioram sn (NY); Quebradillas, 22 Kovember 1913, F. Stevens & W . Hess 5154 ( S Y ) ; Collazo River, 19 February 1926, A?. Britton 8583 ( S Y , S, US); prope Pepino ad Encas, 10 January 1887, P. Sintenis 5827 (F), 5829 (BM, BR, E, G 4 sheets, K, L, LD, MO, NY 2 sheets, PH, P 2 sheets, UCWI, US 2 sheets, TN, 2); ad Encas, 10 October 1887, P. Sintenis sn (GH); Lake Gua- jataca, 25 August 1963, Alain Liogier I0187 (IJ, NY, US); Sierra de Lares in rupibus calcar. ad ?Guajataca,? 3 Febru- ary 1887, P . Sintenis 6096 (C, UCWI, US); Lares, 400-500 m, 6 April 1913, AT. Britton, E . Dritton & W . Hess 2735 (F, NY, US); Lares, 400 m, 31 October 1943, F . Sargent 3248 (US); Lares, 30 June 1914, J . Stevenson & J. Johnston 2060 (US); Lares in praeruptis ad cavernam Pajita, 13 January 1887, P . Sintenis 5845 (BM, C, F, G 3 sheets, GH, GOET, K, L, LD, M, MO, SY 2 sheets, P 2 sheets, S, U, UCWI 3 sheets, US 2 sheets, I$?, Z); Lares in rupibus ad ostium cavernam ?Par- tita,? January 1887, P . Sintenis sn (GH); cliffs along P.R. Hwy 129 about 8 km E of Lares, 27 August 1970, L . Skog 8; J . Skog I707 (BH, US); Esperanza, Dominguito road, Octo- ber 1911, H . Cowles 223 (US); Hato Arriba near Arecibo, 3 March 1914, A?. Britton & J . Cowell 2012 (NY); Hato Arriba, near Arecibo, 1 March 1915, E . Britton 5102 (KY, US); Arecibo, 5 October 1963, T . Talpey I l a (US 2 sheets); near the Arecibo Ionospheric Observatory on P.R. Hwy 625 about 13 km S of Arecibo, 27 August 1970, L . Skog & J . Skog 1715 (BH, US); near radar site, Esperanza, S of Arecibo, February 1965, T. Talpey 27 (BH); Esperanza, 10 October 1963, T . Talpey I l b (US 2 sheets); between Arecibo and Utuado, 4 March 1914, A?. Britton & J . Cou?ell 2042 (?T); Arecibo to Utuado, 14 March 1906, A?. Britton & J . Cowell 1458 ( S Y ) ; between Utuado and Arecibo, 13 March 1906, ,I. Cowell 802 (F, NY, US); Utuado to Arecibo, 14 June-22 July 1901, L . Underwood & R. Griggs 803 (IVY 2 sheets, US 2 sheets); Adjuntas, February 1886, P. Sintenis sn (GH); P.R. Rte 10 north of Utuado, 17 March 1970, M . Meagher 189 (USF); below Utuado, 20 February 1923, N. Britton & E . Britton 7525 (G, GH, NY, US); prope Utuado in sylva N U M B E R 29 111 PUERTO RlCO 50 k m I I I FIGURE 65.-Distribution of Gesneria section Physcophyllon in Puerto Rico. (G. cuneifolia = solid circles; G . ret icdata = solid triangles.) primaeva montis Hueco ad Cayuco in praeruptis calc., 25 March 1887, P. Sintenis 6568 (GH, K); Utuado, in praeruptis ad 10s Angeles, 17 January 1887, P. Sintenis 5935 (K, LD, UCWI); Utuado, 8 November 1913, F. Stevens & W . Hess 4694 (NY); near Florida, 31 January 1925, N . Britton, E . Britton & K . Boynton 8184 (NY); S of Florida, 250 m, 23 August 1963, H. McKee 10617 (P); Manati, 7 November 1913, W . Hess 4137 (NY); Ciales, 21 July 1962, A h i n Liogier 9581 (IJ, NY); Ciales, 24 February 1926, N. Britton 8594 (NY); Ciales, 17 August 1913, J. Johnston 957 (NY); District Baya- mon, Ciales, Bo. Cordillera, 12 December 1937, J. Otero 355 (MO); prope Vega-baya, October 1886, A. Stahl 605b (L); Km 28, San Juan-Arecibo road, 13 January 1935, F. Sargent 131 (US): Toa Alta, 8 February 1927, N . Britton & E. Brit- ton 8791 (NY, s); above Toa Alta, 300 ft, 26 November 1966, T. Talpey 64 (BH); near Toa Aka, toward Esperanza, 26 November 1966, T. TaEpey 63 (BH). LOCALITY UNKNOWN: ?Mexique,? no date, M . Pavon sn [Sesse & Mocino] (G); ?Guadeloupe,? 1846, M . Gamier sn (P); ?Peru,? 1827, M . Pavon sn (P); in rupibus calcareis, August 1827, H . Wydler 453 (G 2 sheets). CULTIVATED: Brooklyn Botanic Garden, 1 May 1923, M . B. sn (BH); Cornell University, April 1960, H . Moore sn (BH), G-285, 18 December 1962, R. Clark sn (BH), November 1956, H. Moore 7172 (BH, US), G-763, 8 October 1964, R. Clark sn (BH), 2 August 1966, L. Leva 38 (BH), G-857, 3 October 1969, L. Skog 1411 (BH), 1 Decem- ber 1965, M. Stone 123 (BH), G-869, 18 March 1969, J. Arnbrose 121 (BH), 14 September 1966, M. Stone 215 (BH, NA); Longwood Gardens, Acc. 5565, 7 December 1960, J. Peek 408 (BH), 31 August 1961, J. Peek 635 (BH). DIscussIoN.-Conradia cuneifolia was described by A. P. de Candolle from the original Sesse and Mocifio drawings, which are now missing. Until these are found, the tracings published by A. de Candolle in 1874 are considered as the types for his new species names. There is a specimen from the Moricand herbarium (now at Geneva) that closely matches the tracing of Gesneria cuneifolia. Although the collector on the sheet is given as Pavon, the specimen was probably collected by Sesse and Mocino in Puerto Rico and later became part of the Pavon herbarium. The authority for the combination Gesneria cuneifolia was determined only after examining all of the available evidence concerning conflicting dates of publication. The combination was pub- lished twice in 1894 and both names could prob- ably be traced back to the original drawings and collections made by Sess.e and Mocino in Puerto Rico during their expedition to Nueva Espana (1787-1804). Stafleu (1967) gives the exact date for the publication of the part of Die naturlichen Pflanzenfamilien containing the new combination Gesneria cuneifolia (A. P. de Candolle) Fritsch as 7 August 1894. The second edition of Flora Mexicana by Sesse and Mocino was published in Mexico also sometime in 1894, but no information has been found to establish the exact month and date of publication, which might prove to be earlier. Until evidence is found that Gesneria cuneifolia of Sesse & Mocino has priority, the combination made by Fritsch is accepted as the earliest. Gesneria cuneifolia has been introduced into cultivation and is widely grown in botanic gardens and as house plants. One collection of seed made 112 SMITHSONIAN CONTRIBUTIONS T O BOTANY FIGURE 66.-Type specimens: a, holotype of Gesneria hybocarpa Urban & Ekman, Ekman H4382 (S); b, lectotype of Gesneria flurflurascens Urban, Wright sn (GOET). (Lectotype is upper plant.) by T. Talpey in 1963 from Quebradillas gorge in Puerto Rico is available under the cultivar name ?Quebradillas.? Other color variants of Gesneria cuneifoliu are grown but have not been given cultivar names. The chromosomes of two culti- vated clones at Cornell University have been counted and found to be n = 28, the highest hap- loid number known in Gesneria. 31. Gesneria hybocarpa Urban & Ekman Gesneria hybocarpa Urban gi Ekman in Urban, Ark. Bot. 22A (lo):%, 1929. Subshrubs: stems woody, decumbent or suberect, up to 20 cm tall or rarely more, bark green, ver- ruculose, and pilose when young, becoming gray, smooth and glabrescent with age; branches from the base, slender, 3-4 mm in diameter at 10 cm below the resinous apex, internodes up to 2 cm long. Leaves alternate, not clustered: petioles sulcate or flattened, 1-6 mm long, ca 1 mm in diameter, green, verrucose and pilose-glandular; blades rhombic to oblanceolate, often falcate, 5.1-14.9 cm long, 1.9-4.2 cm wide, membranous, base cuneate or acute, margin serrulate to strongly serrate, occasionally shallowly undulate, apex acute or acuminate, adaxial surface green, sub- scabrous from remnants of scattered appressed white trichomes, abaxial surface lighter green, NUMBER 29 1 1 3 pilose with sparse appressed white trichomes and scattered glands, veins prominent, verruculose. Inflorescences numerous, each of 1-3 flowers: peduncles terete, up to 3.9 cm long, ca 1 mm in diameter, green, very sparsely pilose; bracts 2, linear-lanceolate, ca 1.5 cm long, ca 1 mm wide, green and pilose; pedicels terete, up to 3.2 cm long, ca 1 mm in diameter, green and pilose; floral tube obconic, 2-3 mm long, ca 2 mm in diameter, pilose and verrucose; calyx spreading, connate for ca 3 mm above apex of ovary, 5-lobed, each lobe narrowly triangular, 1.2-2.1 cm long, ca 2.5 mm wide at base, outside green, pilose with trichomes at margins appressed, veins 3, prominent, inside green and glabrous; corolla tube curved, up to S cm long, ca 2.5 mm wide at base, ca 8 mm broad below the middle and narrowing to ca 4.5 mm ;it the throat, dark red, essentially glabrous with a few trichomes near the limb, limb 5-lobed, each lobe semiorbicular, ca 0.5 mm long, margin erose, mouth oblique; stamens 4, adnate to base of corolla tube and reaching the mouth of the corolla. Stigma exerted 4 mm beyond corolla (vide Urban). Capsule globose, 3-5 mm long, 4-6 mm in diameter, brownish, glabrescent, and verrucose, costae 10; seeds fusiform, twisted, ca 1 mm long, dark brown. TYPE-CoLLECTIoN.-Anse-aaFoleur, Haiti, E. Ek- man H4?82 (S, holotype, Figure 66a; EHH, I J , K, NY, S, US, isotypes). DISTRIBUTION AND EcoLocY.-Gesneria hy bocarpa is known only from northern Haiti (Figure 54), where it grows on limestone cliffs. The type- collection with flowers was made in June. SPECIMENS ExAMINED.-HISPANIOLA. HAITI. Dkpartement du Nord: Massif du Nord, Anse-a-Foleur, at Riviere St. Anne, 21 June 1925, E. Ekman H4382 (S, holotype of Ges- neria hybocarpa Urban & Ekman; EHI-I, IJ, K , NY, S, US 2 sheets, isotypes). DIscussIoN.-Gesneria hybocarpa is a poorly known species based on a single collection by Ek- man (H4382). Only one mature flower was present on the material available, that on the holotype at Stockholm. The apparent sigmoid curve of the red corolla is similar in shape to the corollas of G. acaulis and G. christii. Gesneria hybocarpa differs from both in the prolonged connation of the calyx lobes above the attachment of the corolla to the floral tube, and in the verru- cose petioles and floral tube. 32. Gesneria libanensis Linden ex Morren Gesneria libanensis Linden ex Morren, Ann. SOC. Roy. Agric. Gand 2361, pl. 84, October 1846.-Walp., Repert. Bot. Syst. 6:737, 1847.-W. Hook., Bot. Mag. 74: pl. 4380, 1848.- Fritsch, in Engl. & Prantl, Nat. Pflanzenfam. 4 (3b):184, fig. 81D, 1894.-Urb., Symb. Ant. 2:381, 1901.-Morton in Leon & Alain, F1. de Cuba 4:462, 1957. Rhytidophyllum floribundun Lemaire, F1. Serres Jard. Eur. 2: pl. 178, December 1846 [superfluous name]. Conradia Jloribunda Paxton, Paxton?s Mag. Bot. 15:99, 1848. [Type: plate and description.] Herincquia floribunda (Lemaire) Decaisne ex Herincq, Rev. Hort. ser. 3, 2:323, 1848.-Jacques & Herincq, Man. Gen. P1. 2:562, I850.-Lavallee Horticulteur Francc 1864:177, pl. 11, 1864. Ophianthe libanensis (Linden ex Morren) Hanstein, Lin- naea 26:205, pl. 1: fig. 33, 1854. Conradia corrugata Grisebach, Mem. Amer. Acad. Arts, n.s. 8:526, 1862. [Type-collection: Monte Verde, Cuba, C. Wright 1335 (GOET, holotype; BR, G, GH, K, MO, NY, P, PH, W, isotypes).] Conradia libanensis (Linden ex Morren) Grisebach, F1. Brit. W. Ind. 462, 1862 [pro parte as to Cuban material].- Griseb., Cat. P1. Cub. 201. 1866 [pro parte]. Pentarhaphia libanensis (Linden ex Morren) Hanstein, Lin- naea 34:292, 1865. Pentarhaphia floribunda (Lemaire) Bentham & Hooker f . ex Carriere, Rev. Hort. 1878:30 and plate, 1878.-Anon., Gard. Chron. 1878 (2):593, 1878. Gesneria hondensis Morren ex Jackson, Index Kewensis 1 : 1024, 1893 [sphalm., non Humboldt, Bonpland & Kunth Pentarhaphia corrugata (Grisebach) Gomez de la Maza, Anales SOC. Esp. Hist. Nat. 23:280, 1894. Gesneria libanensis var. corrugata (Grisebach) Urban, Symb. Ant. 2:381, 1901. Gesneria Zopezii Morton, Brittonia 9:19, 1957.-Morton in Leon & Alain, F1. de Cuba 4:462, 1957. [Type-collection: Oriente, Cuba, C. Morton, M . Lopez F . & Alain Liogier 8759 (US, holotype; BM, isotype).] (181 7-1818)l. Slow-growing subshrubs: stems woody, erect or pendent, to 60 cm tall, ca 3 mm in diameter at 10 cm below apex, bark rugose, or smooth, brown to gray, apex pilose, resinous; branches occasional, internodes 0.7 to 2.0 cm long. Leaves alternate in a rosette, deciduous or per- sistent: petioles sulcate, 0.7-2.0 cm long, 1-2 mm wide, green, villous, blades oblanceolate, 1.9-16.0 cm long, 0.6-4.0 cm wide, membranous, base acute, margin crenate to serrate, occasionally irregularly undulate-lobulate, adaxial surface occasionally bullate, or smooth, dark green, glabrous or with appressed pubescence when young, becoming glabrescent and trichomes white with broad bases, 114 ShlITHSONIAN C O N T R I B U T I O S S TO BOTANY veins sunken, abaxial surface lighter green, tri- chomes occasionally reddish, appressed between veins, or glabrous, some veins prominent, pilose, diverging at an angle of ca 45". Inflorescences of 1-5 protandrous flowers, occasionally as long as the leaves: peduncles short, 0.2-2 cm long, sparsely pilose; bracts 2 , linear, u p to 2 cm long, ca 1 mm wide, green, pilose; pedicels 1.5-2.5 cm long, red or green, pilose; floral tube obconic or turbinate, ca 2 mm long, 2.0-3.5 cm wide, green, pilose to villous with reddish or color- less trichomes, costae 10; calyx 5-lobed, lobes con- nate ca 1-2 mm, erect, broadly triangular, 1.5-7.0 mm long, ca 1.0-2.5 mm wide at base of free part, margin entire, apex acute, outside green, sparsely pilose, glandular, reddish or colorless trichomes. becoming white, veins prominent, inside green, glabrous; corolla tube ventricose at the middle from a long attenuate base, 1.9-3.6 cm long, ca 2.5 mm at base, ca 8 mm wide above middle, 3-4 mm wide at throat, outside yellow at base, scarlet above, pilosity more dense toward limb, inside red or yellow, glabrous, mouth oblique, limb somewhat expanded, upper lobes exceeding lower lobes, sub- orbiculate, ca 2-3 mm long, 2-3 mm wide, sparsely pilose, margin dentate, lateral and basal lobes ca 1-2 mm long and wide, margin subentire, sparsely glandular; stamens 4, adnate for less than 1 mm to base of corolla tube, exserted about 2 mm beyond corolla mouth, filaments up to 3.0 cm long, yellow or red, glabrous, anthers oblong, ca 1 mm long, ca 0.5 mm wide, coherent in 2 pairs, staminode ca 3 mm long; inferior ovary pilose at apex with colorless or white trichomes, disc annular, 5-angled, style exserted beyond upper corolla lobes, to 4.0 cm long, yellow or red, glabrous, stigma clavate, recurved, papillate. Capsule ovoid to globose, 3-5 mm long, 3-5 mm wide, green to brown or gray, pilose, dehiscing apically, costae 5 or 10; seeds fusiform, twisted, striated, less than 1 mm long, ca 0.25 mm wide, reddish-brown. TYPE-COLLECTION.-hft. Liban, Oriente, Cuba, J . Linden 1833 (GENT, lectotype, Figure 67; BM, BR, G, K, NY, P, W, isolectotypes). DISTRIBUTION A D EcoLow.-Gesneria libanensis grows in the Province of Oriente in Cuba (Figure 68) on damp limestone rocks and cliffs at 100-700 m elevation. Flowering collections have been made SPECIMENS ExAMINED.-CUBA. PROVINCE OF ORIEKTE: Sierra de Nipe, farallones de Cayo del Rey, 16 April 1940, J. Cara- bia 3527 (XY, US); Cayo Rey, July 1943, Bro. Clemente 2956 (US); Cay0 del Rey, southwestern base of the Sierra de Sipe, ca 100 m, 7 January 1956, C. Morton, Alain Liogier & h.1. Lopez F . 8759 (US, holotype of Gesneria lopezii Morton; BM, isotype); Sierra de Nipe, desfiladero del Rio Bio, 16 September 1922, E . E k m a n 15113 (F, G , K, KY, S, US); Maceo, Cay0 Re), 30 January 1955, M. Lopez F . 1787 (NY); Miranda, Sierra de Xipe, November 1940, Bro. L e o n 19510 (US 2 sheets); Bayate, Picote (in Sierra de Sipe), ca 550 m, 16 July 1916, E . E k m a n 7405 (G , K, SY, S); Bayate in decliv. mont. calcar. Picote, 14 March 1918, E . E k m a n 9148 ( S ) ; Picote in declivibus arduis, 14 November 1919, E. E k m a n 10112 ( S ) : Monte Picote, near Palmito del Cauto, 400 m, 29 January 1936, C . Morton 9669 (US 2, sheets); Florida Blanca, Alto Songo, 450 m, 19 January 1960, Alain Liogier, J. Actna k M . Lopez F . 7402 (US), 5 February 1954, h.1. Lopez F. 1111 (LS), 9 February 1957, M . Lopez F . 2854 (BH, US), 1 May 1957, M . Lopez F . 2873 (US): Represa del Guam, FICCRE 67.-Lectotype specimen of Gesneria libanensis Linden . - - during every month of the year. ex Morren, Linden 1833 (GEKT). NUMBER 29 115 100 k m I FIGURE 68.-Distribution of Gesneria section PhyscophyZlon in Cuba, (G. libanensis = solid squares; G. purpurascens = open circles; G. reticulata = solid triangles.) 28 December 1917, Bros. Hioram & Angel 1510 (LS, NY); Guanahamo, Monte Libanon, San Fernandez, ca 700 m, 24 December 1919, E. Ekman I0282 (G, K , NY, S); on cliffs, Monte Libano, 5 March [1861], C. Wright 3080 (GH); Monte Liban, 1843-1844, J. Linden 1833 (GENT, lectotype o f Gesneria libanensis Linden ex Morren; BM 3 sheets, BR, G, K 2 sheets, NY, P 2 sheets, W 2 sheets, isolectotypes); Monte Verde, January-July 1859, C. Wright 1335 (GOET, holotype of Conradia corrugata Grisebach; GH, isotype), 1859 or 1860, C. Wright 1335 (BR, G 3 sheets, K, NY, P, PH, W); La Perla, 600-650 m, 6-18 February 1911, J. Shafer 8547 (NY); La Alcachofa, Guantanamo, December 1934, Bro. Hioram 8 (LS, NY); Yateras, January 1933, Bro. Hioram 14076 (GH); banks of Rio de Pena, Toa, near Dos Pasos, 400 m, December 1953, AZain Liogier 3647 (GH, NY). LO- CALITY UNKNOWN: Bayate de Monte-Rus, 24-25 February 1955, M. Lopez F . I979 (NY, US); 1861, C. Wright 377 (=3080) (S). DIscussIoN.-Gesneria libanensis Linden ex Mor- ren was probably brought into cultivation by Lin- den, who collected plants from Cuba in 1844 and may have sent living plants or seeds to de Jonghe or Van Houtte (Lavallee, 1864). Within a few years the plant became well known in horticulture and was distributed under many different names. No material attributable to Gesneria libanensis is now known to be in cultivation. Morren (1846) studied both cultivated material and the original collection by Linden, but un- fortunately did not select a holotype. Many dupli- cates of L i n d e n 1833 exist. T h e specimens may not have been collected at the same time, because on the sheet at Vienna is written ?1844 fl. en juin . . .,? while on other sheets flowering is reported as occur- ring in May. The adequate sheet of L i n d e n 1833 at GENT has been chosen as lectotype. According to de Candolle (1880) the Linden collection is at the University of Ghent, but whether this is the main Linden herbarium is doubtful (Van Veken, in litt.). For the application of the name Conradia libanensis sensu Grisebach, see the discussion under Gesneria purpurascens Urban. The type of Conradia corrugata Grisebach, C. W r i g h t 1335, was collected at Monte Verde, a coffee plantation north of Guantanamo. Wright spent many weeks at various times from 1857 through 1861 collecting in the area of Monte Verde (Underwood, 1905), thus, specimens num- bered 1335 were probably collected at different times. Only those dated January-July 1859 can be considered part of the type-collection. 33. Gesneria pedicelluris Alain Gesneria pedicellaris Alain, Mem. New York Bot. Gard. 21 (2):146, 1971. Subshrubs: stems woody, erect or pendent, to 0.75 m long, ca 5 mm in diameter at 10 cm below apex, apex resinous, red or green, becoming brown, bark pilose to densely villous with red articulated 116 SMITHSONIAS CONTRIBUTIOSS T O BOTANY many-celled trichomes, appearing brown in dried specimens, glabrescent below, lenticels obscure; branches usually from the base, internodes usually very short, seldom to 2 cm long. Leaves alternate, usually congested at apex of stem: petioles sulcate, 0.3-1.6 cm long, 1-2 mm wide, usually red, villous with numerous short glands; blades oblanceolate or obovate, falcate, 3.7-16.4 cm long, 1.0-4.2 cm wide, membranous, base cordate, obtuse or cuneate, margin crenate to grossly serrate, apex acute, adaxial surface light or dark green, nitid, scabrous with appressed broad- based white trichomes, veins impressed, abaxial surface lighter green, villous along the red or yellow-green prominent veins, stomata sometimes in groups. Inflorescences axillary, shorter than, equal to, or longer than the subtending leaves, 1- to 4-flowered, erect: peduncles terete, 0.8-10.2 cm long, 1-2 mm in diameter, green or reddish, villous; bracts linear, 1.2-3.0 cm long, ca 1 mm wide, green, pilose, pri- mary pedicels 1.5-2.5 cm long, secondary pedicels 2.5-5.6 cm long, ca 1 mm in diameter, green or red, villous; floral tube obconic, ca 2 mm long, 2.0-3.5 mm wide, yellow, green, or red, villous, 10-ribbed; calyx Nobed, lobes connate for ca 1-2 mm, erect, linear to lanceolate, 0.4-1.4 cm long, 2-3 mm wide at base, outside green, sparsely velutinous, 3-nerved, inside green, glabrous; corolla tube curved slightly above the base and curved (Figure 16f) slightly downward above the inflated middle, sparsely velutinous, 1.9-3.5 cm long, base ca 4 mm in diameter, yellow, middle 5-9 mm in diameter, red-orange, narrowing to 3.5-5.0 mm in diameter at the red throat, becoming dark red on the limb, inside yellow-orange, glandular-pubescent, limb 5-lobed, oblique, lobes erect, orbiculate, erose to serrate, upper lobes ca 2 mm long, 3 mm wide, lateral lobes ca 1 mm long, ca 2 mm wide, basal lobe ca 1.5 mm long, ca 2 mm wide; stamens 4, not exserted at anthesis, filaments linear, curved, slightly shorter than corolla tube, adnate at the base for ca 1 mm, yellow to white, glabrous, except pubescent near middle, anthers oblong, ca 1.5 mm long, ca 1 mm wide, coherent in 2 pairs, pollen yellow, grains isopolar, prolate, size small (22.2 pm long at the polar axis, 10.6 ym wide at the equato- rial axis), tricolpate, colpi 20.6 pm long, apocolpia rounded (Figure 17b), sexine heterobrochate, reticulate, lumina ca 0.3 pm across, muri to 0.5 pm wide; staminode ca 5 mm long; ovary completely inferior, white villous at apex, disc 5-angled, 2 mm wide, white to reddish, glandular, style exserted longer than corolla tube, curved, up to 3.5 cm long, white, pubescent near base, stigma clavate, papillate. Capsule globose, persistent calyx lobes re- flexed, 3-4 mm long, 4-5 mm wide, brown, pilose to glabrescent, costae 10, dehiscent at the apex; seeds striated, ca 0.5 mm long, 0.2 mm wide, tawny, nitid. TYPE-COLLECTION.-Near Abreu, Dominican Re- public, Alain Liogier 16150 (NY, holotype, Fig- ure 69; F, GH, IJ, P, US, isotypes). DISTRIBUTION AND EcoLocu.-Gesneria pedicel- laris is known only from the north coast of Do- minican Republic (Figure 54), where it grows on shaded steep limestone cliffs at elevations from near sea level to ca 50 m. Flowering collections have been made of this species in the wild during January to March, July, and September. In the greenhouse plants bear flowers throughout the year. SPECIMEKS ExAMI~ED.-HISPANIOLA. DOMINICAN REPUB- LIC. Province of Maria Trinidad Shnchez: ca 2 km TY of Abreu, ca 50 m, 8 July 1970, L. Skog 1589 (BH, US 2 sheets); near Abreu, about sea level, 28 September 1969, Alain Lio- qier 16150 ( S Y , holotype of Gesneria pedicellaris Alain, consisting of two sheets; F 2 sheets, US, isotypes); near Cabrera, ca 100 m , 28 September 1969, Alain Liogier 16154 ( S Y , US); El Diamante, Cabrera, about sea level, 21-23 Jan- uary 1972, Alain Liogier I N 6 7 (NY); near la playa del Diamante, between Julia Molina and Cabrera, 1 March 1953, J . Jimenez 2563 (BH, E, RDJ, US); between La Entrada and Abreu, 19 February 1965, T . Ta lpey 33 (BH, E, US); El Diamante, north from Sagna, 19 February 1965, T . Talpey k J. Jirnenez 5120 (BH, RDJ, US), T . T a l p e y 5120 ( S Y ) . CULTIVATED: Cornell University, G-898, 5 October 1969, L. Skog 1415 (BH), 23 June 1966, M . Stone 132 (BH); G-1226, 5 October 1969, L. Skog 1409 (BH), 3 November 1970, L. Skog 1749 (BH); G-1395, 14 March 1972, L . Skog 1819 (BH). DIscussroN.-Gesneria pedicellaris (introduced by T. Talpey in 1965) was available in the horti- cultural trade long before it was described as a new species. The plants, however, were distributed as either G. christii Urban, native to southern Haiti, or G. barahonensis Urban, from southern Dominican Republic, both of which G. pedicel- laris resembles. Gesneria pedicellaris can be dis- tinguished from both by the scabrous leaves and the bracts more than 1.2 cm long, from G. barahonensis by the corolla tube more than 1.9 NUMBER 29 117 FIGURE 69.-Holotype specimens of Gesneria pedicellaris Alain, A h i n Liogier 16150 (NY). cm long, and from G. christii by the erect orienta- tion of the flowers in living material and by the leaf margin, which is crenate or serrate, not lobulate. A few hybrids have been produced with species not closely related to Gesneria pedicellaris (see Table 7 ) . Backcrosses to the parents have not been successful (Table 8). The hybrid plants (G-1231, G-1239, G-1245, G-1420) appear intermediate in most morphological characters, but due to their large size, none of the hybrids are likely to have horticultural potential. 34. Gesneria purpurascens Urban Gesneria purpurascens Urban, Symb. Ant. 2:380, 1901.-Mor- ton in Leon and Alain, F1. de Cuba 4:461, 1957. Gesneria yumuriensis N. Britton & P. Wilson, Mem. Torrey Bot. Club 16: 109, 1920 [?yamuriensis?].-Morton in Leon and Alain, F1. de Cuba 4:462, 1957. [Type-collection: Near Rio Yamuri, Cuba, J. Shafer 7786 (NY, holotype; US, isotype).] Slowgrowing subshrubs: stems woody, erect or pendent to ca 20 cm tall, 1 cm in diameter at 10 cm below apex, bark smooth and villous to rugose and glabrescent, green or reddish above, becoming brown to gray, apex slightly resinous; branches few, internodes to 0.9 cm long. Leaves alternate, usually closely spaced in a rosette: petioles subsulcate, 0.3-2.0 cm long, 1-2 mm wide, green or reddish, villous with reddish or tawny trichomes; blades spathulate or narrowly obovate, broadest beyond the middle, seldom falcate, 3.9-21.9 cm long, 1.8-5.0 cm wide, mem- 118 SMITHSOXIAX COXTRIBUTIONS TO BOTASY branous, base cuneate to cordate, margin crenate, apex acute or obtuse, adaxial surface strongly bullate, dark green, glandular pubescent when young, glabrescent with age, trichome bases en- larged and white, lateral veins diverging from the midvein at an angle usually greater than 45O, abaxial surface lighter green, with appressed white hairs between veins, veins prominent, trichomes on veins red or green. Inflorescences of several flowers, about one-fifth the length of the subtending leaves: peduncles very short, ca 5 mm long, bracts linear, ca 1.5 cm long, ca 1 mm wide, reddish, pilose, pedicels 0.8- 2.5 cm long, reddish, pilose to villous; floral tube obconic or turbinate ca 2 mm long, 2-3 mm wide, reddish, villous, costae obscure; calyx lobes 5, erect, connate 1-2 mm at base, each narrowly triangular, 0.4-1 cm long, ca 2 mm wide at base of free part, margin entire, apex acute, curved, out- side red, sparsely pilose, 3-nerved, inside red, gla- brous, but glandular; corolla tube ventricose at middle from a narrow base, 2.8-3.3 (-4.0) cm long, 2-3 mm in diameter at base, broadening to ca 9 mm above middle, then narrowing to 2.5-4.0 mm at throat, outside yellow at base with red lines, scarlet above, more densely pilose toward apex, inside yellow or with red lines, glabrous, mouth oblique, limb slightly expanded, lobes orbiculate, usually suberose to erose, rarely subentire, upper lobes ca 2-4 mm long, ca 3 mm wide, lateral lobes ca 1.5 mm long, ca 2 mm wide, basal lobe ca 1 mm long, ca 2 mm wide, dark red, glandular, becoming stipitate-glandular at the margin; stamens 4, shortly adnate to base of corolla tube, barely ex- serted beyond mouth to ca 2 mm, filaments ca 3 cm long, yellow, glabrous, anthers oblong, 1.0- 1.5 mm long, 0.5-1.0 mm wide, yellow, coherent in pairs, staminode ca 6 mm long; ovary inferior, apex pilose, disc annular, 5-angled, style exserted, ca 3.2 cm long, curved at apex, yellow or reddish, sparsely pubescent at base, stigma clavate, papillate. Capsule broadly turbinate, ca 3-4 mm long, ca 6 mm wide, gray-brown, pilose or glabrescent, dehiscing at the apex, costae 5 or 10; seeds fusi- form, striate, ca 1 mm long, ca 0.5 mm wide. TYPE-COLLECTION.-oriente, Cuba [ 1860-1 8641, C . W r i g h t 3080 (BM, G, K, MO, P); Baracoa, Oriente, Cuba [1861], C. W r i g h t , s n (GOET, lectotype; Figure 66b). DISTRIBUTION AND EcoLoGY.-Gesneria purpur- ascens is known from near the northern coast of the Province of Oriente in Cuba (Figure 68), where it grows in moist places on limestone cliffs from near sea level to 400 m elevation. Flowering specimens have been collected in November through April and August. SPECIMENS ExAMINED.-CUBA. PROVINCE OF ORIENTE: northern slope of El Yunque de Baracoa, 300 m, 2 January 1960, Alain Liogier & M. Lopez F . 7178 (US), 14 January 1960, Alain Liogier & J. Acuna 7533 (US): near the base of El Yunque de Baracoa, toward the east, ca 300 m, 14 January 1960, Alain Liogier & J. Acuna 7541 (US), 7546 (US); El Yunque de Baracoa, 300 m , 14 January 1960, A h i n Liogier & J . Acuna 7696 (US); side and top of El Yunque, 20 De- cember 1910, J. Shafer i 8 9 f (XU); on rocks by Rio Maca- guanigua, 20 March 1931, L. Bailey I5153 (BH); Baracoa ad Rio Macaguanigua, 19 January 1915, E. Ekman 4328 (G, K , SY, S); Baracoa, March 1903, L. Underwood & F . Earle 235 (SY); vicinity of Baracoa, 18 February, 11 March 1910, J. .Shafer 3916 (F, NY 2 sheets, US); Santiago, vicinity of Bara- coa, 24-29 January 1902, C. Pollard, E . Palmer & W . Palmer 42 (CV, F, GH, MO, NY, P H , US); Mata, Baracoa, 14 Janu- ary 1956, Alain Liogier 8: C. Morton 5101 (GH); Bahia d e Mata, Baracoa, 1 February 1952, J . Actcna & Diaz Barreto 17379 (SV, US); ad Mata Bay, 26 Kovember 1914, E. Ekman 3653 (SY, S ) ; Abra del Yumuri, Baracoa, Alain Liogier & C. Morton 5092 ([13 April 19561 G H , [13 January 19561 IJ); Rio Yumuri, entre Baracoa y Maisi, 26 August 1939, Bro. ? Leon 17249 (US 2 sheets); cliffs near the mouth of the Rio Yumuri, east of Baracoa, near sea level, 13 January 1956, C. Morton & Alain Liogier 9023 (BM, US 2 sheets), 9038 (BM, US); Yumuri prope Baracoa, 25 November 1914, E . Eknzan 3636 (F, G, K , S, US); near Rio Yamuri (sic), 6, 8 December 1910, J . Shafer 7786 (NY, holotype of Gesneria yumuriensis N. Britton & P. 1Vilson; US, isotype); Baracoa, 1861, C. Wright sn (GOET, lectotype of Gesneria purpuras- cens Urban); [locality unknown] 1860-1864, C. Wright 3080 (BM, G 3 sheets, G H , K, MO, P, isolectotypes). DIscvssIor\..-The typification of Gesneria pzir- pzirascens Urban is complex and probably in- volves four taxa. In 1846, Morren described Gesneria libanensis, ostensibly from cultivated material, but cited L i n d e n 1833 from Monte Liban, Cuba, as the type. Grisebach, in 1862, made a new combination, Conradia libanensis, based on hiorren?s description, but misapplied the name to Jamaican and Haitian plants citing col- lections by MacFadyen and Lane (now at Kew) from Jamaica. In 1866, Grisebach applied the name ?Conradia libanensis to L i n d e n 1833 and W r i g h t 3080. Although, C . W r i g h t 3080, found in many herbaria, is usually a mixed collection, one NUMBER 29 119 part is truly Gesneria libanensis Linden ex Morren. In 1901, Urban realized the disparity between the Wright collection and the species described as Gesneria libanensis. Urban?s solution follows: (1) the misapplied 1862 name for the Jamaican plants he placed under a new variety of Gesneria acaulis Linnaeus; (2) he cited Grisebach?s 1866 use of Conradia libanensis under Gesneria l iban- ensis Linden ex Morren, excluding the 1862 interpretation; and (3) he described a new species, Gesneria purpurascens, based on that part of W r i g h t 3080 collected in 1861 at Baracoa. Later Urban (1912), in an observation under Gesneria barahonensis Urban, cited Conradia libanensis sensu Grisebach as to the Haitian plants men- tioned by Grisebach in 1862, but wrote that he had not seen the Grisebach material. Subsequent search for material from Hispaniola annotated by Grisebach as Conradia libanensis has not been successful. The specimen Urban designated as the type was probably at Berlin and is no longer extant. No other material of W r i g h t 3080 from Baracoa, dated 1861, has been found. Wright apparently numbered his specimens by taxon rather than by collection, and he rarely supplied collection data on the sheets. Only one Wright collection of Ges- neria purpurascens, from Baracoa, 1861, has been found annotated by Urban, but unfortunately it bears no collection number. In view of Wright?s haphazard numbering system, this specimen (at Gottingen) was very likely part of the type gather- ing and is here selected as lectotype to stabilize the nomenclature. The type of Gesneria yumuriensis N. Britton & P. Wilson, J . Shajer 7786, differs from typical specimens of Gesneria purpurascens? only in being more glabrescent and resinous. In all other char- acters the species are identical. Specimens of both species have been collected from the type-locality of Gesneria yumuriensis and show a range in pilosi ty. 35. Gesneria reticdata (Grisebach) Urban Gesneria reticulata (Grisebach) Urban, Symb. Ant. 1:478, Conradia reticulata Grisebach, Cat. P1. Cub. 201, 1866. Conradia reticulata 0 obovata Grisebach, Cat. PI. Cub. 201, 1866. [Type-collection: Yunque de Baracoa, C. Wright sn (GOET, holotype).] 1900 & 2:378, 1901 [pro parte, as to basionym]. Pentarhaphia reticulata (Grisebach) Gomez de la Maza, Anales SOC. Esp. Hist. Nat. 23280, 1894. Pentarhaphia reticulata 0 obovata (Grisebach) Gomez de la Maza, Anales SOC. Esp. Hist. Nat. 23:280, 1894. Gesneria cuneifolia var. obouata (Grisebach) Morton, Brit- tonia 9:19, 1957. Gesneria cuneifolia var. disjuncta Morton, Brittonia 9:19, 1957. [Type-collection: Rio San Juan, Dominican Repub- lic, G . Miller 1244 (US, holotype).] Subshrubs: stems herbaceous or seldom woody, erect, to 20 cm tall, 2-4 mm in diameter at 5 cm below apex, green or reddish, glandular-resinous at apex, becoming brown to gray below; branches from near the base, internodes to 1.6 cm long, leaf scars prominent above. Leaves alternate, rosulate or clustered at apex or spread along rapidly growing stems: petioles sulcate, broad, 1-3 mm long, ca 2 mm wide, green or reddish, glandular-pubescent or glabrous; blades oblanceolate, ovate, obovate or spathulate, 1.8- 11.4 cm long, 0.7-5.2 cm wide, membranous, smooth or bullate and wrinkled especially at the midvein, base cordate to cuneate, margin entire to serrulate to grossly dentate, often ciliate, apex acute, rounded or rarely emarginate, adaxial sur- face light to dark green, glabrous or with sparse appressed trichomes, rarely pilose toward the base, abaxial surface lighter green or reddish, glabrous or with sparse appressed pubescence, veins prom- inent and strongly arching, stomata scattered. Inflorescences cymose, of 1-3 flowers, usually ca two-thirds the length of the subtending leaves: peduncles terete, 1-7 mm long, usually short, ca 1 mm in diameter, green or red, glandular; bracts linear or oblanceolate, 3-7 mm long, ca 1-2 mm wide, green or red, glabrous; margin entire or serrate, pedicels terete, 1.5-5.4 cm long, ca 1 mm in diameter, green or red, glabrous; floral tube broadly obconic, 2-3 mm long, 4-7 mm wide, green or red, glabrous, alate; calyx 5-lobed, porrect to patent, lobes connate ca 1.5 mm at the base, ovate, 4-7 mm long, ca 2 mm wide at base, 2-4 mm wide at middle, apex acute, margin entire, occasionally ciliate, outside green or reddish, gla- brous or sparsely pilose, with 3 prominent veins, inside green or reddish, glabrous; corolla tube cylindric, with a slight bend at throat, tube 1.1-2.7 cm long, 2.5-3.2 mm wide at base, not gibbous, 2.5-3.5 mm at middle, not ventricose, 3.5-4.5 mm in diameter at throat, outside red, orange-red, or 120 SMITHSONIAN CONTRIBUTIONS T O BOTANY pallid yellow, pilose, inside yellow to orange, gla- brous, limb flaring to patent, 1.0-2.1 cm wide, sparsely pubescent, lobes 5, serrate or notched, upper lobes obovate, 5-6 mm long, 6-7 mm wide, lateral lobes 5-6 mm long, 4-6 mm wide, basal lobe 4-6 mm long, 5-8 mm wide; stamens 4, adnate for less than 1 mm to base of corolla tube, not ex- serted, filaments slightly curved, 0.9-2.4 cm long, reddish, yellow, or white, glabrous, anthers oblong, ca 1.5 mm long, 1 mm wide, coherent in two pairs; ovary completely inferior, apex pilose, disc 5- angled, unlobed, yellow, glabrous, style curved, 1.0-1.8 cm long, reddish, yellow to white, glabrous, stigma clavate. FIGURE 70.-Type specimens: a, isotype of Conradia reticulata Grisebach and Gesneria reticulata (Grisebach) Urban, Wright 308I, including label bearing number 3081 (GH); b, isotype of Conradia reticulata fl obovata Grisebach (= Gesneria Capsule erect, broad to globose, ca 3-5 mm long, 4-8 mm wide, brown to gray, glabrous, de- hiscing at the apex, costae prominent, persistent sepals patent; seeds broadly fusiform, twisted, ca 0.5 mm long, 0.25 mm wide, reddish-brown to black, nitid. TYPE-CoLLECTIoN.-Yunque de Baracoa, Cuba, C. Wright 3081 (GOET, holotype; BM, G, GH (Figure 70a), K, isotypes). DISTRIBUTION A D EcoLocu.-of all known spe- cies of Gesneria, G. reticulata is the most wide- spread with a center of distribution in the moun- tains of Hispaniola and outliers in the Oriente Province of Cuba and in the Luquillo Mountains of eastern Puerto Rico (Figures 54, 65, and 68). I t s habitat is usually wet woods, near or at the edge of streams and waterfalls on igneous or lime- stone rocks from 80 to 1200 meters elevation. This species has been collected with flowers in all months except September. In cultivation, plants may flower continuously. Bay, 5 December 1914, E. Ekinan 3753 ( S ) : on the road to Qu ib iah , Baracoa, 80 m, 1 January 1960, AEain Liogier & M . Lopez F . 7151 (US): N of El Yunque, Baracoa, 150 m, 13 January 1960, Alain Liogier & J. Acuna 7609 (US): inner valley of El Yunque, Baracoa, ca 500 m, 14 January 1960, Alain Liogier & J. Acuna 7543 (US); a t the base of El Yunque de Baracoa, 200 m, 2 January 1960, Alain Liogier & M . Lopez F . 7247 (US); El Yunque in rupibus calcareis, 17- 18 December 1914, E. Ekman 3912 ( S ) ; slopes and summit of El Yunque, near Baracoa, 1000-2000 ft, 30-31 January 1902, C . Pollard & W . Palmer I37 (CU, F , GH, MO, NY, PH, US): El Yunque Mt., Baracoa, March 1903, L . Under- wood & F . Earle 622 (NY), 690 (NY), 1062 (NY); Yunque de Baracoa, June [1861], C. Wright sn (GOET, holotype of Conradia reticulata fi obovata Grisebach; GH, isotype); Arroyo Frio, Sra. de Imias, So. Baracoa, 950 m, 17 J u l y 4 August 1924, Bro. Leon I2180 (GH, NY, US). LOCALITY UNKNOWN: ?Cuba,? 1860-1864, C . Wright 3081 (GOET, holotype of Conradia reticulata Grisebach; BM, G, GH, K, isotypes). HISPANIOLA. HAITI. Departement du Sud: Morne de la Hotte, ca 800 m, 10 June 1917, E . Ekman HI32 ( C , S ) ; Riviere Glace, 750 m, 6 August 1945, L. Holdridge 2184 (US); Massif de la Hotte, group Morne Rochelois, Mira- goane, on the path Lebrun to Quatre-Chemins, ca 700 m, 30 March 1927, E . Ekman H7923 ( S ) ; Departement du Nord: Massif du Nord, Port-de-Paix, top of Haut Piton, 1100-1205 m, 6 April 1925, E. Ekman H3711 (EHH, S, US); vicinity of St. Louis du Nord, 30 March-7 April 1929, E . Leonard & G . Leonard 14245 (GH, K , MO, NY, US), 2 April 1929, E . Leonard & G . Leonard 14248 (US) , G April 1929, E. I.eonard & G . Leonard 14472 (US); Massif du Sord, Baveux, top of SPECIMENS ExAMINED.-CUBA. PROVINCE OF ORIENTE: Taco ret icdata) , Wright sn (GH). Morne Brigand, 900-1 150 ?m, 20-21 December 1924, E. E k k n SUMBER 29 121 H2972 ( S ) ; Port Margot to Correil, 1200-1500 ft, 7 August 1903, G. Nash 196 ( N Y ) ; ravine NW of Marmelade, 2230 ft, 3 August 1905, G . Nash 8: N. Taylor 1353 (NY). Departe- ment de 1?0uest: Morne Fourmi, 900 m, June 1905, W. Buch 765 ( I J ) ; Massif de la Selle, Croix-des-Bouquets, Badeau, ca 1200 m, 14 March 1927, E. Ekman H7851 (A, S ) ; Massif de la Selle, Port-au-Prince, Morne Malanga, ca 1200 m, 27 January 1926, E. Ekman H5457 (EHH, F, G, GH, IJ, NY, S, US), DOMINICAN REPUBLIC. Province of Pedernales: Agua Negra, Sierra de Bahoruco, 23 March 1967, E. Marcano 5274 (NY, RDJ); Agua Negra, 750 m, 22 March 1967, T . Talpey 77 (BH); Agua Negra, Sierra del Baoruco, 800 m, 17 February 1969, Alain Liogier 13949 (NY). Province of Bara- hona: Caniada Maluca, above Barahona, 27 July 1952, R. Howard 12192 (A, US); 1912, M . Fuertes 1111b ( K ) . Prov- ince of Monte Cristi: Distr. of Sabaneta, bank of Rio Cidra, 5-600 m, 30 November 1930, E. Valeur 569 (US). Province of Santiago: overlooking Rio Antonzape Malo, Mata Grande, San Jose de las Matas, 1000 m, 1-7 Octo- ber 1968, Alain Liogier 12965 (NY); h T O y 0 del Toro, Tamboril, 20 February 1965, E . Marcano & J . Jime?nez 5121 (E, NY, RDJ, US); Arroyo del Toro, February 1965, T . Tal - pey 35 (BH). Province of La Vega: Casabito, 6 April 1968, E . Marcano & T . Talpey 539f (NY, RDJ); Casabito, 10 April 1968, E . Marcano & T . Talpey 5428 (NY, US 2 sheets): 22 km E of El Rio along road toward the Santiago-Sto. Domin- go Hwy, 900 m, 12 August 1970, G. Davidse 2660 (BH, MO); road from Bonao to Constanza, 1000 ft, 29 May 1970, M . Meagher 295 (USF); ca 20 km from El Rio near road from Bonao to El Rio, 16 July 1970, L. Skog 1606 (BH, US 2 sheets); Piedra Blanca, 10 December 1947, H . Allard 17795 (US); 19 January 1948, H. Allard 18902 (BH, US); near Piedra Blanca, at km 73 of the Carretera Duarte, ca 200 m, 12 February 1929, E. Ekman H11544 ( K , S , US); banks of Maimon river, 13 April 1954, J. Jime?nez 2607 (E, US); near Rio Maimon, along the road from Piedra Blanca to Maimon, ca 500 m, 6 July 1970, L. Skog 1586 (BH, US). Province of Puerto Plata: Gurabito de Yaroa, coffee plantations and woods, Yaroa valley, 350-500 m, 1 May 1968, Alain Liogier 11028 (NY); Cordillera de Yaroa, limestone ridge facing Yaroa valley, 800-850 m, 11 May 1968, Alain Liogier 11234 (F, NY, USF). Province of Duarte: Quita Espuela, vicinity of San Francisco de Macoris, 400-1000 m, 5-17 April 1922, W. Abbott 2087 ( I J , NY, US). Provinces of Espaillat or Maria Trinidad Sanchez: Rio San Juan, 22 March 1928, G. Miller 1241 (US, holotype of Gesneria cuneifolia var. dis- jztncta Morton). Province of San Cristobal: Cabirma de la Loma, 600 m, 28 November 1970, Alain Liogier 17752 (NY): Cabirma de la Loma, 19 March 1967, E. Marcano 5207 (BH, S Y , RDJ); Cabirma de la Loma, NW of San Cristobal, 2000 ft , 9 April 1968, T . Talpey 92 (BH); Los Guineos, San Cris- toba1, 24 March 1958, E . Marcano & J . Jimener 3656 (RDJ, US). Province of Samana: Yuna river, 28 February 1963, Bro. Basilio Augusto 736 ( N Y ) . Province of El Seibo: Liali, banks of Rio Lajiagua, 100-500 m, 8-20 February 1923, W . Abbott 2623 (NU, US). LOCALITY UNKNOWN: ?St. Domingo,? 1857, R . Schornburgk sn (K). PUERTO RICO: Sierra de Luquillo, in monte Jimbnes, 17 July 1885, P. Sintenis 1343 (BM, G 3 sheets, GH, GOET, K, L, LD, M, P, S, UCWI, US); El Yunque, 6 October 1963, T . Talpey 12 (US); El Yunque forest, km 10 on Route 191, 12 February 1965, T . Talpey 25 (BH, U S ) ; at km 10 on Palmer to Florida road through Luquillo Mts., 15 August 1963, R. Wagner 353 (A): on the sides of the waterfall at marker 10.2 km on P.R. Hwy 191, in the Caribbean National Forest (Luquillo Experi- mental Forest), 500 m, 5 June 1970, L. Skog 1517 (BH), 20 August 1970, L. Skog & J . Skog 16f4 (BH); km 14 of Route 191 through Luquillo Mts., 3 April 1965, R. Howard & L. Nevling 15775 (A, US) . CULTIVATED: Cornell University, G-784, 7 October 1964, R . Clark sn (BH), 3 October 1969, L. Skog 1412 (BH), 16 February 1966, M . Stone 117 (BH, ?4); Cornell University, G-897, 5 October 1969, L. Skog 1413 (BH), 22 December 1967, M . Stone 350 (BH). DIscussIoN.-Plants of Gesneria reticulata have been brought from Puerto Rico and the Domini- can Republic and grown at Cornell University under the accession numbers G-784 and G-897, respectively. The plants from Puerto Rico avail- able in the horticultural trade under the name Gesneria czineifolia ?El Yunque? were first col- lected at El Yunque peak in the Luquillo Moun- tains by Talpey in 1963 (Talpey, 1966). As a cultivar ?El Yunque? was first validly published in 1966 by Katzenberger (as Gesneria cunefolia var. ?el Yunque? [sic]). ?El Yunque? has been errone- ously misapplied (Elbert, 1973) to plants in culti- vation but probably originally collected by McKee from near Florida in central Puerto Rico. The latter plants are part of Gesneria cuneifolia, while cv. El Yunque correctly refers to cultivated mate- rial of Gesneria reticulata. Section 7. Dittuntheru (G. Don) L. Skog, new transfer Conradia section Dittanthera G . Don, Gen. Syst. 4:650, 1838. Pentarhaphia section Codonoraphia (Orsted) Bentham in Bentham & Hooker, Gen. PI. 2:1005, 1876. Pentarhaphia section Vaupellia (Grisebach) Bentham in Bentham & Hooker, Gen. P1. 2:1005, 1876. Gesneria section Codonoraphia (Orsted) Fritsch in Engler & Prantl, Sa t . Pflanzenfam. 4 (3b):184, 1894. Shrubs or trees: stems erect, to 8 m tall; branches from the base and from the upper leaf axils. Leaves lanceolate, elliptic to obovate, occasion- ally falcate or oblanceolate, membranous to sub- coriaceous, plane to subbullate, glossy. Inflorescences 1- to 4-flowered, exceeding the subtending leaves; corollas infundibuliform-cam- panulate, green, yellow, reddish-brown, or rose; 122 SMITHSOSIAN CONTRIBUTIOSS T O BOTANY stamens long-exserted, or extending only to the corolla mouth. Capsule turbinate or elongate, costae prominent. D1scussros.-The ancestral stock of section Dit- tanthem probably included plants similar to those now found in section Pentaj-haphia. From this ancestral source of fruticose, hummingbird- pollinated plants, Dittanthel-a may hake eiolved as a response to bat-pollination mechanisms. In habit and vegetative characters, this section re- sembles sections Pentarhaphia, Lachnoblaste, and lily J vi ek inn t h e. The species in section Dittanthel-a appear to be distinct from each other b) combinations of char- acters and are separated geographically. Any one of the characters may be found in more than one of the species, but not alwajs in the same combina- tions. The large number of taxa in this section have probably evolved due to the high concentra- tion of possible pollinators in the small land area in Jamaica and the other islands. T o stabilize the concept of section Dittantlzera, Gesnel-ia exserta, one of the species included in Don?s circumscription, is here selected as lectotjpe species. LECTOTYPE-SPECIES.-GeSneJ ia eXSel-ta STVartZ. 36. Gesneria aZpina (Urban) Urban Gesneria alpina (Urban) Urban, SJmb. Ant. 5:498, 1908.- Gesnerin cal>cosa Tar. alpinn Urban, Sjmb. Ant. 2:3?6, 1901. Shrubs: stems woody, spreading, bark rugose, graybrown, to 6 m tall, verrucose, lenticels elon- gated; branches 5 mm in diameter at 30 cm from the apex, lower branches to 6.1 m long (fide Stearn), nodes enlarged, leaf scars prominent. Leaves alternate or approximate: petioles sul- cate, 0.7-1.5 cm long, 1-2 mm wide, green to brown, glabrou5, verrucose to smooth; blades ovate to obovate, occasionally falcate, 6.0-1 1.4 cm long, 3.5-4.7 cm wide, membranous, base acute or rounded, margin 5errate to dentate, apex acuminate, adaxial surface dark green, glabrous, abaxial surface lighter green, glabrous, veins prominent, verrucose. Inflorescence I-flowered, in leaf axils near branch apices: peduncles terete, 1.0-1.6 cni long, 1-2 mm in diameter, glabrous, resinous; bracts 2 , linear, 4 mni long, less than 1 mm wide, brownish, resinous; pedicels 3.1-3.9 cm long, 1-2 mm in Adams, F1. P1. Jamaica 681, 1972. diameter, wider toward the apex, green, glabrous; floral tube turbinate, ca 4 mm long, 7 mm in diameter, glabrous, resinous; calyx lobes 5, con- nate for less than 1 mm at base, each lobe sulcate at base, 1.3-1.9 cm long, 2-3 inm wide at the base, outside dark green, glabrous, resinous, inside pilose, resinous;. corolla campanulate, tube 1.3-2.3 cm long, 6 inm wide at the base, expanding to ca 2 cin \vide at mouth, both sides yellow, glabrous, resinous, limb 5-lobed, each lobe glandular, upper lobes ca 8 inm long, 6 mm wide, margin dentate, lateral and basal lobes ca 8 mm long, 9 mm wide, margin subentire; stamens 4, shortly adnate to corolla base, not exserted, filaments linear, ca 16 inm long, pale yelloiv, glabrous, glandular, anthers oblong, 2 min long, 1 mm wide, red or brown on back, glandular; ovary inferior, disc annular, 4 mm long, yellow, puberulent with glandular trichomes, style ca 2.3 cm long, exserted, green, puberulous, glandular, stigma stomatomorphic. Capsule turbinate, dehiscing into four valves, ca 1.0 cm long, 8 mm wide, green to gray, glabrous, costae 5; seeds fusiform, twisted, ca 1 mm long, red to brown (Figure 19m). TYPE-CoLLECTIos.-Near Blue YIoun tain Peak, Jamaica, W . Harris 7547 (BM, lectotype, Figure f l u ; GH, K, NY, UCWT, isolectotypes). DISTRIBLTION A D EcoLocu.-Gesneria alpina is known only from the parishes of St. Andrew and St. Thomas in eastern Jamaica (Figure 72) at high elevations of 1100-2200 m. Its habitat is usu- ally limestone soil in deeply shaded, \vet, moun- tain forests. Gesneria alpina appears to flower only during the winter months, November to March. The corolla could accommodate the heads of the larger flower-feeding bats in Jamaica, Biachyphylla piimila and Phyllonycteris aphylla, and probably the heads of smaller bats as well, all of which may be pollinators. summit cone of John Crow Peak, Blue Mountains, 5500-5700 ft, 24 Sovember 1954, G. Proctor 9926 (IJ); Vicinity of Cinchona, Sew Haven Gap, 2-10 September 1906, A?. Britton 150 ( S Y ) ; Cinchona, Blue Mountains, leeward slopes, 9 February 1915, J . Harris R: J . Laurence GI9129 (US); Cin- chona, Blue Mountains, windward slopes, 2 March 1915, J. Harris Rc J . Larc~rence C1532f ( S Y ) ; Cinchona, Blue Moun- tains, leeward slopes, 9 March, 191.5, J . Harris R: J . Louq- rence Cl i39f (US) Cinchona, Blue Mountains, windward slopes, 13 March 1915, J . Harris R: J . Lawrence C15470 (F): Blue Mts., near Abbey Green, 3300 f t , A . Rehder $72 (A). PARISH OF ST. T H O M ~ S : above Portland Gap, 5800 f t , 18 SEPCIhfEXs E X . ~ ~ ~ I ~ \ ? E O . - - J A ~ I A I C A . PARISH OF ST. A4iYDREW: NUMBER 29 Key to the Species of Section Dittanthera 1. Caljx lobes connate in a cup for o w r half their length above the o\ar); floral tube and capsule linear, elongate, and slender; Jamaica 37. G. calycina 1. Caljx lobes connate foi less than one-tenth of their length abole the ocar); floral tube and capsule nariowly tuibinate to nearl) spherical. 2. Stamens and st)le long exserted, tvice as long ds the corolla tube 01 more. 3. Calyx lobes suipassing corolla, sulcate at base: inflorescences 1-flowered: capsules 1 5 cm long or more; Jamaica 38. G. calycosa 3. Calyx lobes not exceeding coiolla, nor sulcate; inflorescences of (1-) 2 or more f l o ~ - ers; capsule less than 1 cm long 4. Leaf bases acute, calyx lobes mole than 6 mm long; Puerto Rico 4. Leaf bases truncate to cordate; cal5x lobes i mm long oi less; Jamaica 43. G. $edunculosa 40. G. exserta 2. Stamens and stjle less exseited, or included, one and one-half times a$ long as the corolla tube 01 less. 5 Inflorescences of t vo or more floneis; corolla green with puiple macalae; margin of corolla limb fimbriate; Hispaniola 41. G. fruticosa 5. Inflorescences of a single floxei corolla dull rose, gieen to )ellow \\ith no maculae; margin of corolla limb dentate to subentiie: Jamaica. 6 Caljx lobes flat a t base; apex of stem, petioles and inflorescences pilose, rarelb resinou5 39 G . clundestina G CalJx lobes sulcate at base; apex of stem, etc , glabrous, iesinous 7. CalJx lobes suipassing cotolla, spntred on innei surface abo\e sulcus, apex awned, corolla green, reddish-brow n or dull rose, tube 1 .l-1.3 cm long 42 G . onychocalyx, ne\$ species 7. Calyx lobes shorter than or seldom equaling corolla, not spurred abo\e sulcus, apex not obviousl) awned; corolla yellow, tube 1 3-25 cm long 36. G . aZ$ina 123 February 1962, C. Adams 10700 (UCTVI); Portland Gap, ca 5250 f t , 13 December 1994, G. Proctor 9616 (IJ, XY, US): Portland Gap, ca 5290 f t , 8 January 1956, G. Proctor Br W . Steam 11451 (IJ); near Blue Mountain Peak, 6500 ft, 30 Sovember 1898, W . Harris 7547 (Bhi, lectotype of Gesneria calycosa var. alpina Urban; GH, I(, SY, UCTYI, isolecto- types); near Blue hit. Peak, 31 January 1893, 14?. Harris sn (Bhf, UC\VI); Blue hit. Peak, 13 December 1890, A . Hitch- cork siz (F, h I 0 ) ; west slope of Blue Mt, Peak, 6000-7000 f t , 8 Sovember 1954, G. Proctor 9 f i 5 (A, IJ, NY); by side of trail, west spur of Blue Mt. Peak, 6000-6500 ft, 8 January 1956, It?. Stearn 100 (A, BH, BM, I(, P, S, CCTVI). LOCALITY UNKSOWS: ?America Meridionalis,? no date, Shakespear sn (BM). DIscussrox.-Since no authentic holotype speci- men has been discovered, the specimen of Harris 7547 at British l luseum (Natural History) has been selected as lectotype. 37. Gesneria calycina Swartz Gestzeria calycii~a Swartz, Prodr. 90, 1788.-Sw., F1. Ind. Occid. 2:1026, 1800 (?Gemera?).-Lunan, Hort. Jam. 1:322, 1814.--Spreng. Sjst. Veg. ed. 16, 2:839, 1825; Fritsch in Engler k Prantl, Kat. Pflanzenfam. 4 (3b):184, 1894.-Urb., Symb. Ant. 2:376, 1901 .-.4dams, F1. P1. Jamaica 680, 1972. Conradia ralycina (Swartz) Xiartius, 9ov. Gen. Pi Sp. 3:38, 1829 Pi 3:191, 1832.-G. Don, Gen. Sp t . 4:651, 1838.-DC., Prodr. 7:929, 1839. Sinningia calycina (Swartz) Hort. ex Don in Loudon, Hort. Brit. 241, 1830. Gloxinia calycina (Sirartz) Hort. ex Steudel, Nom. Bot. ed. 2, 1:690, 1840 (?calicyna?). Gloxinia l indleyi Steudel, Xom. Bot. ed. 2, 1:690, 1840 [pro parte, as to Swam synonym]. Paupellin calycina (Swartz) Grisebach, F1. Brit. TV. Ind. 460, 1862. Pentarhaphie calycina (Swartz) Hanstein, Linnaea 34:307, 1865. Shrubs or trees: stems woody, erect, bark ex- foliating, gray-brown, lenticels numerous, erum- pent, subverrucose, resin abundant apically and covering younger leaves; branches from enlarged nodes, green-brown, glabrous, internode length less than 1.5 cin. Leaves a1 ternate to approximate, crowded at branch apices: petioles sulcate, 1 .O-3.O cm long, 1-2 mm wide, green, glabro~s, verrucose; blades 124 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 71.-Type specimens: a, lectotype of Gesneria calycosa par. a l p h a Urban and Gesneria alpina (Urban) Urban, Harris 7547 (BM); b, holotype of Conradia clandestina Grisebach and Gesneria clandestina (Grisebach) Urban, Wilson sn (K). elliptic to oblanceolate or obovate, 12.0-21.6 cm long, 2.8-5.5 cm wide, membranous to subcori- aceous, base acute to cuneate, margin slightly crenate to dentate, apex acute to caudate, both sides green, glabrous, abaxial veins prominent, Inflorescences axillary, 2- to 4-flowered: pedun- cles terete, 5.8-11.5 cm long, 1-2 mm in diameter, green to brown, verrucose; bracts 2, linear to lanceolate, 0.8-2.0 cm long, 1-4 mm wide, cadu- cous, glabrous, apex acute; pedicels terete, 1-4 cm long, 1 mm in diameter; floral tube linear, 5-8 mm long, elongating in fruit, 1-2 mm wide, green, glabrous, verrucose, resinous; calyx lobes 5, con- nate at the base for over half their length into a cup, each free portion flat, membranous, ovate, ' resinous occasionally. 0.6-1.6 cm long, 4-8 mm wide, apex acute to rounded, margin entire, green and glabrous both sides, veins prominent; corolla obliquely sub- campanulate, tube 0.8-1.2 cm long, ca 1.6 cm wide at mouth, inside and outside green, glabrous, limb 5-lobed, each lobe erect, green, rounded at apex, margin subentire; stamens 4, adnate to the base of the corolla tube, exceeding the corolla tube by 6 mm, filaments linear, 2.0-2.5 cm long, greenish, glabrous, anthers oblong, ca 2 mm long, 1 mm wide, not coherent; ovary inferior, disc annular, entire, style linear, broader toward apex, ca 2 cm long, greenish, glabrous, stigma stomatomorphic. Capsule elongate, splitting into 2 valves from the apex, 1.2-2.0 cm long, 4-5 mm wide, gray-brown, glabrous, verrucose, costae 10, prominent; seeds S U M B E R 29 125 FIGURE 72.-Distribntion of Gesneria section Dittanthera in Jamaica. (G. alpina = solid squares; G. calfciiia = open squares; G. calycosa = solid circles; G. clandestina = open triangles; G. exserta = open circles; G. otifchocalyx = solid triangles.) linear, twisted, 1 mm long or slightly longer, less than 0.25 mm wide, reddish-brown (Figure 19f). TYPE-CoLLEcTIoh?.-?India occid: Jamaica,? 0. Sruartz s n ( S , holotype, Figure 73a; BM, LINN, M, UPS, isotypes). DISTRIBUTIOS AND EcoLocY.-Gesneria cdyc ina is found in the John Crow Mountains of eastern Jamaica (Figure 72), on moist shaded limestone slopes at 400-500 m altitude. Gesneria calycina has been collected with flowers from November to April. This species may be pollinated by any of the flower-feeding bats in Jamaica (Table 6). SPEcIifEsS ExAwNED.-JAMAICA. Parish of Portland: 5 mi SIV of Priestman?s River, ca 1500 ft, 6 February 1953, G. Proctor 7635 (IJ); Ecclesdown, 1200 ft, 1 March 1961, C. A d a m 9096 (UCIVI 2 sheets), 29 March 1961, C. Adams 9343 (BXI, UCIVI); John Crow Mts. above Ecclesdown, 1350 ft, 8 April 1967, A . Katzenberger & R. Katzenberger 108 (RDJ); east slope of John Crow Mts., 1.5 mi SIV of Eccles- down, 1500 ft, 3 April 1931, G. Proctor 5682 (IJ, U S ) ; east slope of the John Crow hits., 1.5-2 mi SIV of Ecclesdown, 1500-2000 ft, 14 January 1955, G. Proctor 9800 (IJ); east slope of the John Crow &Its., 1-1.5 mi SIV of Ecclesdown, 1 March 1961, G. Proctor 22125 (BM, GH, IJ, US); SFY of Ecclesdown, lG00 ft, 3 Sovember 1964, T. Talpey 13 (BH); 1-2 mi SIV of Ecclesdown, John Crow hits. foothills, 1900- 2500 ft, 30 March 1958, T. Yuncker 18541 (BM, F, G, I J , S); Betty?s Hope, near Manchioneal, 3000 ft, 20 March 1858, A?. Ll ? i kO?? 463 (GOET, K ) . LOCALITY U N R N O M . ~ : ?Jamaica,? no datc, P i ? . Alacfadyen stz (K) ; ?India occid: Jamaica,? n o date 0. Su?artz sn ( S , holotype of Gesneria calycina Swartz; Bhi, L I S S , hi, UPS, istoypes). DIscussIoN.-Gesneria caZycina is unusual in the genus in having the calyx lobes connate for over half their length into a cup above the apes of the ovary. T h e distinctiveness of the very striking calyx was realized by Grisebach (1859-1864) when he erected a new genus, Vaupellia, for the species. T h e connation in Gesnel-ia calpcina is an elabora- tion of the usual condition in the genus, calyx lobes connate for 1-2 mm at the base. I n other characters this species resembles others in section Dit tan thera. 38. Gesneria calycosa (W. Hooker) 0. Kuntze Gesneria calycosn ( IV . Hooker) 0. Kuntze, Rev. Gen. 473, 1891 (?Gesnera?).-Fritsch in Engl. & Prantl, S a t . Pflan- zenfam. 4 (3b):184, fig. El.%, 1894.-Urb., Symb. Ant. 2:376, 1901.--Adams, F1. P1. Jamaica 681, 1972. Conradia calycosa 11?. Hooker, Icon. P1. 7: sub pl. 689-690, 1844.-IValp., Repert. Bot. Syst. 63395, 1847. Pentarhaphia calycosa (1%?. Hooker) Decaisne, Ann. Sci. S a t . Bot. scr. 3, 6:99, 1846.--T%?alp., Repert. not. Syst. 6 : 7 3 5 , 1847.-Hanst., Linnaea 26:214, fig. 32.4, 1834.-Griseb., F1. Brit. I V . Ind . 460, 1862.-Hanst., Linnaea 34:304, 1865. Codonoraphia cnlycosa (I%?. Hooker) grsted, Cent. Gesn. 68, 1858. Shrubs or trees: stems woody, erect, to 5 m tall, becoming whitish to graybrown, lenticels ca 2 mm long, white, apex resinous, pith reddish; branches many, internodes 0.5-2.0 cm long, nodes swollen. Leaves alternate: petioles sulcate, 1.5-2.4 cm long, 1-2 mm wide, green, glabrous, resinous when young; blades elliptic to oblong, 9.0-17.5 cm 126 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 73.-Type specimens: a, holotype of Gesneria calycina Swartz, Swartz sn (S); b, holotype of Conradia calycosa Hooker and Gesneria calycosa (Hooker) 0. Kuntze, Purdie 635 (K). long, 3.8-6.2 cm wide, membranous, base acute, margin subentire to serrate-crenate, apex acumi- nate, adaxial surface green, glabrous, abaxial side lighter green, glabrous, with a rusty appearance from flaking resin, veins prominent. Inflorescences from axils of young leaves, I-flowered: peduncles slightly flattened, 1.0-2.8 cm long, ca 2.5 mm in diameter, glandular, verrucose; bracts 2, linear, 0.5-2.0 mm long, less than 1 mm wide; pedicels slightly flattened, 2.8-4.7 cm long, ca 2 mm in diameter, green, glandular; floral tube turbinate, 6-8 mm long, 0.6-1.1 mm wide at apex, glabrous, resinous; calyx lobes 5, connate at the base for 1.5 mm above ovary apex, each lobe erect, lanceolate, sulcate at base, 2.6-4.4 cm long, 3-5 mm wide at base, both sides green, glabrous, resinous, midvein prominent; corolla obliquely subcampanulate, tube 1.6-2.0 cm long, 1.4-1.5 cm wide at the mouth, outside yellow-green to cream, glabrous, resinous, inside greenish-white, glabrous, limb 5-lobed, patent to suberect, ca 2.5 cm broad, each lobe semiorbiculate, ca 8 mm long, 8 mm wide, margin erose, glandular; stamens 4, adnate to base of corolla tube for about 1.5 mm, exserted about 2 cm beyond corolla mouth, filaments linear, narrowing to apex, curved, to 4 cm long, 1.5 mm wide, green, glabrous, anthers rectangular, 3-7 mm long, 2-3 mm wide, green to pale brown, coherent in 2 pairs by their apices, pollen grains isopolar (Figure lSj), with a nearly circular amb, tricolpate, prolate, size medium (25.5 pm long at the polar axis, 11.0-14.0 pm wide at the equatorial NUMBER 29 127 axis), sexine reticulate, heterobrochate with larger lumina near the equator, 0.5-1.0 pm wide, at the poles less than 0.25 pm wide, colpi long and narrow, staminode 6 mm long, bearing rudimentary anther; ovary inferior, green, glabrous at apex, disc 5- lobed, green, resinous, style linear, becoming wider near apex, to 4.5 cm long, 1 mm in diameter, green, glabrous, stigma stomatomorphic. Capsule turbinate, elongate to clavate, splitting from the apex almost to the base into 2 or 4 valves, smooth to verrucose, gray-brown, costae 10, prom- inent; seeds linear, twisted, 1-2 mm long, 0.25 mm wide, tawny to brown (Figure 19g). TYPE-CoLLEcTIoN.--Sedburgh, Jamaica, W . PUY- d ie 635 (K, holotype, Figure 7 3 b ; GOET, isotype). grows in the Cockpit country of central to western Jamaica (Figure 72), in damp forests among lime- stone hills. Gesneria calycosa appears to flower during every month of the year; in the greenhouse this species has flowered continually for two years. The pollinator is probably the flower-feeding bat, Phyllonycteris aphylla. DISTRIBUTION AND EcoLocY.-Gesneria calycosa SPECIMENS ExAMINED.-JAMAICA. PARISH OF HANOVER: interior summit slopes of Dolphin Head, 1500-1700 ft, 11 April 1955, G. Proctor 10024 (IJ, NY); interior summit slopes of Dolphin Head, 1500-1700 ft, 7 March 1960, G. Proctor 20688 (IJ). PARISH OF WESTMORELAND: 1 mi WR?W of Hope- well, 1750 ft, 21 November 1955, G. Proctor 11228 (IJ); vicinity o f Woodstock, 24 October 1960, G. Proctor 21517 (BM, GH, IJ); 0.5 mi SCV of Darliston, 1300-1500 ft, 24 February 1970, G. Proctor 31232 (IJ); Clarks Wood district, SE of Woodstock, 1300 ft, 24 April 1961, K. Kramer 1718 (U, W A G ) ; Clarks Wood district, SE of TVoodstock, 1200- 1400 ft, 6 February 1961, G. Proctor & W. Mullings 22001 (BM 2 sheets, GH, IJ) Clarks Woods, S of Exeter, 1300 ft, 11 April 1967, A . Katrenberger & R. Katzenberger 112 (RDJ). PARISH OF ST. JAMES: Mocho Road, Catadupa, 1900 ft, 14 February 1906, W . Harris 9178 ( F , R?Y 2 sheets, UCWI, US). PARISH OF ST. ELIZABETH: Near Maggotty, 950 ft, 28 Decem- ber 1961, C. Adams 10197 ( B M , M O , UCWI). PARISH OF TRELAWNY: Windsor estate, ca 400 ft, 23 August 1956, G. Proctor 15684 ( B M , F, GH, IJ); Windsor Cave, 5/6/1966, I. Cornman sn (UCWI); Windsor, 100-150 m, 1 April 1931, G. Miller 1479 (US); ca 3.5 mi TVNW of Quick Step near Paynes Patent, 9 March 1949, C. Lewis sn (IJ, US). PARISH OF MASCHESTER: Inglewood to Endeavour near Mile Gully, 1750 ft , 23 December 1968, C. Adams 13097 (UCWI); vicin- ity of Mandeville, 15-26 February 1910, S. Brown 205 ( N Y , PH, US); vicinity of Mandeville, 26-30 April 1910, J. Craw- ford 662 (PH, US), 668 (XY); Sedburgh, December 1843, W . Purdie 635 ( K , holotype of Conradia calycosa W. Hooker; GOET, isotype). PARISH OF ST. ANN: Albion Pen, 2000 ft, 12/5/1915, W. Harris 12004 ( B M , F, K , MO, NY, GCWI, US); 1.2 mi E of Albion, ca I500 ft , 4 July 1965, G. Webster 13627 (US). LOCALITY UNKNOWN: Port Royal Mountains, June 1843, W. Purdie sn (K); ?Jamaica,? 1844, W . Purdie sn (K); ?Jamaica,? no date, W . Purdie s n (K, TCD); Jamaica, no date, W . Wright sn (BM). DrscussroN.-GGesneria calycosa differs from all other known species of Gesneria in having in- florescences of one flower with stamens more than twice as long as the campanulate, green corolla. 39. Gesneria clandestina (Grisebach) Urban Gesneria clandestina (Grisebach) Urban, Symb. Ant. 2:377, Conradia clandestina Grisebach, F1. Brit. W. Ind. 461, 1862. Pentarha@& clandestina (Grisebach) Fawcett, Prov. List, 1901.-Adams, F1. PI. Jamaica 681, 1972. Jam. 28, 1893. Shrubs: stems woody to within 30 cm of the apex, to 5 m tall, bark rugose, gray-brown, glabres- cent, lenticels elongated, white, erumpent, becom- ing verrucose with age; branches from base and upper leaf axils, to 1.0 cm in diameter, green to red, pilose with reddish-articulate hairs, primary internodes to 12 cm long, others much shorter, 0.2-1.0 cm long. Leaves alternate: petioles sulcate, 0.8-2.7 cm long, 2-4 mm wide, reddish, pilose with reddish- articulate trichomes; blade elliptic to oblong or oblanceolate, often falcate, 6.9-14.7 cm long, 3.1- 6.1 cm wide, membranous, base rounded to cune- ate, margin serrate or dentate, ciliate, apex acu- minate to caudate, adaxial surface dark green, sparsely pilose, glossy, pubescent, abaxial surface lighter green, pilose along the prominent veins. Inflorescences of one flower: peduncles terete, 4-5 mm long, 1-2 mm in diameter, red, pilose; bracts 2, lanceolate, about 2 mm long, less than 1 mm wide, red, pilose; pedicels terete, 0.9-1.8 cm long, elongating in fruit, 1-2 mm in diameter; floral tube turbinate, ca 6 mm long, 7 mm wide at apex, red, densely pilose; calyx lobes 5 , connate for less than 1 mm at the base, each lobe erect, narrowly ovate, 2.2-2.7 cm long, 5 mm wide above the base, margin entire, ciliate, apex narrowly acuminate or caudate, outer surface reddish, densely appressed-pilose, midvein prominent, in- ner surface reddish-green, sparsely pilose; corolla broadly campanulate, tube 1.7-2.5 cm long, 6-8 mm wide at the base, gradually wider to the 128 ShlITHSONIAN C O N T R I B U T I O S S T O BOTANY mouth, 1.5-2.5 cm wide, outside greenish-yellow, densely puberulent, inside green, glabrous, limb 5-lobed, each lobe subpatent, broadly ovate, glan- dular, margin entire, upper lobes ca 8 mm long, 9 mm wide, lateral lobes ca 8 nim long, 7-11 mm wide, basal lobes 6-8 mm long, 8 mm wide; sta- mens 4, adnate to the base of the corolla tube, exserted for about 4 mm beyond the corolla mouth, filaments linear, ca 2.9 cm long, red, sparsely pilose, anthers oblong, 4 mm long, 2 mm wide, coherent in 2 pairs by their apices, pollen grains (Figure 18a) isopolar, with a nearly circular amb, size small (18.7-19.2 ym long at the polar axis, 11.6-13.7 pm wide at the equatorial axis), tricolpate, colpi 16 pin long, 1 ym wide, apocolpia nearly truncate, prolate, sexine appearing homobrochate, lumina 0.25 ym across, muri about 0.5 ym across, verrucose, warts 1.25 pm across, scattered at ran- dom, about 1 pm apart; ovary inferior, disc annu- lar, style about 3.5 cm long, reddish pilose, stigma stomatomorphic. Capsule turbinate, 1.0-1.3 cm long, 0.9-1.1 cm wide, red, glabrescent, costae of 5 prominent wings; seeds narrowly fusiform to linear, less than 1 mm long, red to brown. TYPE-CoLLECTIoN.-?~amaica,? N . Wilson sn (K, holotype, Figure 71b; GOET, K, isotypes). DISTRIBUTION A D EcoLocY.-Gesneria clandes- tina grows in shaded wet mountain forests in eastern Jamaica (Figure $2) at elevations of 350- 725 m. This species has been collected in flower in January, March, and June, with fruits collected about two months later; plants have not yet flowered in cultivation. A s with Gesneria alpina, this species may be visited and pollinated by Brachyphylla pumila and Phyllonycteris apkylla among the flower-feeding bats. SPECIMESS EX,~~fINED.-J.4M.4IC,~. PARISH OF ST. T H O M A S : Cuna Cuna Gap and vicinity, 600-725 m, 13-19 June 1926, W. A f a x o n 9153 (XY, US); south slopes of Corn Puss Gap, 1600-2000 ft, 1 January 1967, G. Proctor 27755 (IJ); vicinity of Corn Puss Gap, ca 1800 ft, 1 January 1965, G . Gastony 113 (GH): south slopes of Corn Puss Gap, ca 2000 ft, 29 January 1967, G . Proctor 27785 (GH, IJ, U, US); south slope of Corn Puss Gap, 1 March 1969, G. Proctor 29991 (IJ): on the trail to Corn Puss Gap, about 500 f t . from the gap, and 6 miles from Bath, 11 .4ugust 1970, L. Skog 1623 (BH. UCIVI); John Crow Mountains, valley, SE foothills, 350 m , 2-4 March 1909, S. Brittoti 39 f I ( S Y ) ; south eastern foot- hills of John Crow Mts., 2 March 1909, I+?. Harris 8: S. Brittoti 10679 (F , SY, UCWI). LOCALITY U i w ~ o w s : ?Ja- maica,? no date, A?. Wilson sn (K, holotype of Conradia rlandestina Grisebach; G O E T , K, isotypes). DIscussIoN.-Gesneria clandestina is probably most closely related to G. nlpina and G. onycko- calyx; however, both differ from this species in being glabrous and in other characters. Gesneria alpina has a yellow corolla and is more spreading in its growth habit. Gesneiia onychocalyx can be distinguished by its longer flower stalks, which are up to 4 cm long, and by its spurred calyx lobes. 40. Gesneria exserta Swartz Gesneria exserta Swartz, Prodr. 90, 1788.-Sw., F1. Ind. Occid. 2:1024, 1800 (?Gesnern?).-Lunan, Hort . Jam. 1:322, 1814.--Spreng., Syst. T?eg. ed. 16, 2:839, 1825.-Urb., Symb. .4nt. 2:369, 1901.-.4dams, F1. P1. Jamaica 681, 1972. Conradia exserta (Swartz) Martius, Sov. Gen. & Sp. 3:38, 1829 & 3:191, 1832.-G. Don, Gen. Syst. 43650, 1838.-.4. P. tle Candolle, Prodr. 7 : 5 2 5 , 1839. Gesneria rorynibosa Herbarium Balbis 8: Bertero ex .4. P. de Candolle, Prodr. 7 : 5 2 5 , 1839 [non Swartz (1788), nom. nud . pro syn.]. Pentnrhaphin catalpif lorn Decaisne, Ann. Sci. Nat. Bot. ser. 3 , 6:100, 1846 (?catalpaeflora?).-IValp., Repert. Bot. Syst. 6:733, 184i.-Griseb., F1. Brit. IT. Ind . 460, 1862.-Hanst., Linnaea 34:303, 1865. [Type-collection: ?Jamaica,? J . Wiles s n (G, holotype).] Pentarhaphin lessertiana Decaisne, .4nn. Sci. S a t . Bot. ser. 3 , 6:101, 1846.--Walp., Repert. Bot. Syst. 6:735, 1847.- Hanst., Linnaea 34:302, 1865. [Type-collection: ?Jamaica, high mountains,? J . W i l e s s n (G, holotype).] Pentarhapliia parvipora Decaisne, Ann. Sci. S a t . Bot. ser. 3, 6:102, 1846.-IValp., Repert. Bot. Syst. 6:735, 1847.- Hanst., Linnaea 34:302. 1865. [Type-collection: ?Hb. Deles- sert,? (collector unknown), (G, holotype).] Gesneria quadripom Herbarium Lambert ex Decaisne, Ann. Sci. Nat. Bot. ser. 3, 6:102, 1846 [nom. nud. pro syn.]. Pentarhaphia exserta (Swartz) Decaisne, Ann. Sci. S a t . Bot. ser. 3 , 6:104, 1846 [pro parte, as to the synonyms of Mar- tius and Swartz and the description by Swartz]. Codonoraphin catalpiflorn (Decaisne) Orsted, Cent. Gesn. 68, 1858 (?catalpaeflora?). Codonornphin lessertinnn (Decaisne) Orsted, Cent. Gem. 68, 1858. Codonorapliin p a n i f i o r a (Decaisne) Ors ted , Cent. Gesn. 68, 1858. Gesnerin rntnlpif lorn (Decaisne) 0. Kuntze, Rev. Gen. 2:453, 1891 (?Gesnern?). Gesneria lessertiana (Decaisne) 0. Kuntze, Rev. Gen. 2:473, 189 1 (?Gesn ern? ) , Gesnerin pnruiporn (Decaisnc) 0. Kuntze, Rev. Gen. 2:473, 1891 (?Gesnern?). Shrubs or trees: stems woody, erect, to 5 in tall, to 2 cm in diameter, bark reddish-brown, resinous NUMBER 29 129 near apex; branches many, 5 mm in diameter, reddish-brown, glabrous and resinous, lenticels elongated, erumpent, internodes 0.3-3.5 cm long, shorter toward the apices, nodes slightly swollen. Leaves alternate: petioles sulcate, 1.5-3.5 cm long, 2-3 mm wide, green to brown, glabrous, resinous, not verrucose; blades ovate, elliptic to obovate, 6.3-13.5 cm long, 2.0-5.3 cm wide, sub- coriaceous, plane, base subcordate to truncate, oblique, margin crenate, apex acute to acuminate, adaxial surface dark green, glabrous, glossy, veins immersed, abaxial surface lighter green, glabrous with prominent veins. Inflorescences 1- to 4-flowered: peduncles terete, 4.1-6.0 cm long, 1-2 mm in diameter, reddish- brown, glabrous, glandular-resinous; bracts 2, lanceolate, 1-2 mm long, less than 1 mm wide, very resinous; pedicels terete, 1.3-2.8 cm long, 2 mm in diameter, green, glabrous, resinous, with elongated lenticels; floral tube ca 5 mm long, 7 mm wide, green, glandular-resinous; calyx lobes 5, each linear-lanceolate to triangular, 4-5 mm long, 3 mm wide at the base, apex acute, withering at anthesis, green and very resinous, with one prominent vein; corolla narrowly campanulate, tube ca 1.5 cm long, gibbous at the base, then contracted to 7 mm in diameter, then becoming wider at the mouth, ca 12 mm broad vertically, 7 mm wide horizontally, both inner and outer surfaces of the tube greenish white, glandular, margin suberose and ciliate, upper lobes semiorbiculate, ca 6 mm long, 7 mm wide, erect to patent, lateral lobes ca 9 mm long, FIGURE 74.-Type specimens: a, holotype of Gesneria exserta Swartz, Swart% sn (S); b, holotype of Gesneria onychocalyx L. Skog, new species, Proctor 9799 (BM). 130 SXfITHSOXI,4X COSTRIBUTIOSS TO BOTANY 10 mm wide, reflexed, lower lobe ca 6 mm long, 6 mm wide, reflexed; stamens 4, adnate to the base of the corolla, exserted at anthesis ca 2 cm beyond the mouth of the corolla tube, filaments linear, straight, ca 3.5 cm long, 1 mm in diameter, green, glabrous, anthers rotund, 1 mm long, 1 mm wide, green, glabrous, coherent in two pairs by their apices, staminode 5 nim long, lacking fertile anther; ovary inferior, apex glabrous, disc 5 - angled, ca 5 mm long, 3 mm wide, yellow-green, style linear, to 4 cm long, 1 mm in diameter, green, glabrous but for a few appressed trichomes at the base, stigma stomatomorphic, papillate, nectar abundant, clear and viscid. Capsule nearly spherical, dehiscing from the apex into 2-4. valves, 5-6 mm long, 6 mm in diam- eter, I>rownish, glabrous, resinous, costae 10, not prominent; seeds linear, twisted, about 1 mm long, less than 0.5 mm wide, reddish-brown. TYPE-COLLECTIoS.-?India occidentalis: Jamai- ca,? 0. Swartz sn (S, holotype, Figure 740; G-DC, LINN, M, UPS, isotypes). CHROMOSOME SUMBER.-R = 14 (Lee, 1966a). DISTRIBUTIOS AND EcoLocu.-Gesneria exsevta is endemic to Jamaica (Figure 72) growing in thickets on steep, rocky, shaley slopes or on wooded limestone hillsides at altitudes from 75 to 1200 m. There are collection records for this spe- cies from every month except May. Flowering speci- mens could not be found in Jamaica in early August despite observation in many localities. In the greenhouse, G. cxscrta has produced flowers in December, January, and April. Bat pollination seems likely for G. exserta, possibly by Glossophaga so rir in a a n t ill a r 11 ?n . SPECIMENS EX4sflSED.-JAh~..iIC1Z. PARISH OF ST. ELIZA- BETH: Malvern to Mountainside, 1600 ft, 27 March 1962, C . A d a m 10919 (110, LTCT$?1); Santa Cruz Mountains, Bide- ford district, southwest of Malvern, 1000-2000 ft, 12 August 1954, G. Webs ter & G. Proctor 5317 (A, BM 2 sheets, US); Bideford district, 1.5 mi d u e S of Malvern, 2000 ft, 25 Janu- ary 1964, G. Proctor 2-1513 (BM, MO). PARISH OF CLARES- DON: Broi\.n?s Gate to Thomson Town, Mocho hfountains, 2000 ft, 26 June 1960, C. A d a m 7283 (MO, UCTVI); 2 miles southeast of Crooked River PO, 700 ft, 11 April 1953, G. Proctor 7870 (IJ, SY). PARISH OF Sr. CATHERISE: Bog TValk, 250 ft, 6 December 1953, G. Proctor 8287 (IJ, US, USF); betuwen Flat Bridge and Kensington along Cobre River, 6 Sovember 1964, T . T a l p e y 22 (BH). PARISH OF ST. MARY: Castleton Road, 600 ft, 27 February 1896, E . Cnmpbe l l 6246 (UCIVI). PARISH OF ST. ANDRE!?: Swain Spring near Coopers Hill, 1900 ft, 29 August 1965, B . Morley 83 (UCTVI); Coop- ers Hill, Red Hills, 21 March 1954, G. Proctor 8496 (IJ, XY): Red Hills, on way to Coopers Hill, on rocky slope mi ESE of summit, 2000 ft, 1 January 1956, 1V. Stearn 62 (A, BH, BM 2 sheets); Golden Spring, 267 m, 14 August 1900, TV. Fau*cett 7986 (F, SY); Golden Spring, 800 ft, 14 August 1900, W. Thompson 7986 (UCTVI); Hermitage Dam, ca 1750 ft, 8 January 1944, A . Barry sn (IJ); Moresham River, west side, ?/4 mi above Hermitage Dam, 7 March 1956, TV. .$learn f l l (A, BH, BM 2 sheets); along road between hlt. James and Aft. Airy, ca 1500 ft, 31 January 1970, G. Proctor 31212 (IJ); Road to Gordon Town, February 1885, J . Hart 1136 (UCTVI): Bellevue, 4500 ft, 27 January 1888, H . Eggers 3 i M (B, L, US): Flamstead, January 1927, I. Maxwel l sn (BM); along trail between Gallorvay Gap and Flamstead, Port Royal Mountains, ca 3000 ft, 13 December 1947, A . van d e r Porten s n (IJ); Bellevue, near Flamstead, April 1952, X. Robbins 1514 (UCTL?I); near Bellevue, 3000-4000 ft, 17 Soveniher 1957, T . Ytincker 1 i f O f (BM, F); Guava Ridge, ca 3000 ft, 26 January 1955, G. Proctor 9811 (A, I J , SY, US); hit. Rosanna, 3800-4000 ft, 23 October 1964, A . Skel - ding sti (UCTVI); ca 1 m i A? of TVoodford o n path to Hard- war Gap, 2.500 ft, 14 February 1957, E . Robertson 3220 (UCTI?I 2 sheets); Irish Town, 2450 f t , 20 December 1959, C. A d a m $ 5821 (Bhl, CCTVI); vicinity of Kewcastle, Irish Town, 11 September 1908, A?. Britton 3f52 (XY); along road to Newcastlc about 2 mi above Irish Town, 12 August 1970, L. Skog 1630 (BH): road to Content, 2500 ft, 20 December 1905, W. HaIris 912f (BM 2 sheets, F 2 sheets, K, S Y , UCTVI, 1:s); Charlottenburg, 7 March 1960, A . Skelding 6705 (UCTVI): ?Big Rock,? below Cinchona, 11 March 1920, W . M n x o n & E . Kirrip 976 (A, F, G H , NY, US); near Green River, 011 trail from Cinchona to TVhitfield Hal l , 22 March 1920, W. A i ~ x o n 8; E . Killip 1351 (A , BM, F, H G , SY, US); B ~ L W hlountains. banks of Green River, 2500 ft, 13 February 1903, A . Rehder 517 (A 2 sheets, KY): Clydesdale, 3,300 ft, 7 Sovember 1948, R. Bengry sn (IJ, US); Sorbrook, 600 ft, 27 December 3895, E. Campbell 6136 (NY, UCTVI, US); S o r - hrook, 600 f t , 27 December 1895, W?. Harris 6136 (F); Hal- berstadt, Port Royal Mountains, 2500 It, 1924, C. Norman 188 (Bhf): Halls Delight, 500 m , 15 January 1895, If?. Harris 5 5 f i (BM, F. SY, UCTVI, US): bei Kingston in d. anderen Bergcn, Decembcr 1881, F . Lelimanri 982 (Bhi); Sky Ridge ahove Penfield, 2000 f t , 18 January 1959, A . Skeld ing 5150 (I(). P.lorton, C., .\lain Liogier, k hl . I,ope7 F., 8759(32) Sash , G , 196(35) Nash, G., k N. Tailor, 1353(35) Sicolson, D., 2065(18b) Norman, C., 188(40) Osmaston, H., 5004(20b); 5176(46c) Otero, J., 356(30); 422(43); 618(44c); 771(44c) Palmer, TV., & J. Riley, 153(25): 520(25) Parker, C., sn(l8a) Pavon Herbarium, SIl(30); sn(43) Peek, J., 635(30); 408(30) Perkins, J., 487a(27a); 1025(27a); 1256(45a) I?errottet, G., 230(18b); sn(l8b) Pee, A., 409(43); 866(43) Poeppig, E., sn(25) Pollard, C., & TV. Palmer, 119(47b); 137(35) Pollard, C., E. Palmer 82 Ti7. Palmer, 42(34); 263(47b) Ponthieu, de, sn(l8b) Porten, .4. van der, sn(1a); sn(40) Poulter, J., in .Adam 10895(46d) Poulter, J., et al., & C. Adams, 10893(20d) Powell, I)., 140(27a): 1642(27a): 1669(20b) Prior, R. (earlier Alexander), 558(40); 559(40); 560(27a) Proctor, G., 4007(46d); 4131(46d); 4896(20b); 5616(46d): 5682 (37): 5731(46c); 5732(46c); 6428(27a): 6614(27a); 6926(27a); 7273(20b): 7300(46d): 7635(37): 7684(40); 7870(40); 8148(27a): 8213(46b); 8287(40); 8496(40): 944.5(36); 9526(36); 9527(46a); 9619(36): WO(9); 9779(46b); 9799(42); 9800(37); 9801(46d); 9841(40); 10024(38); 10050(20b); 10205(la); 11228(38); 11424 (40); 11732(12); 11781(40); 15684(38): 15756(20d): 16357(25): 16403(29): 16641(20b): 17856(18b); 19228(18b): 20688(38): 20749(46b); 20845(20c); 20856(20d); 21443(27a); 21517(38); 21750(18b): 21862(46b): 22125(37): 22145(27a); 22160(27a): 22238(20c); 22762(27a); 22965(40); 22974(9); 22991(46d): 23137(20h): 23313(40); 23762(46a); 24271(27a): 24429(40): 24493(27a); 24513(40): 24731(46d): 2475.5(9): 25618(12): 25623(9); 25912(18a); 26320(9): 26617(27b): 26688(20b): 26844(40): 27755i39): 27785(39); 28559(27a); 29306(27a); 29329(27a); 29991(19); 29992(46d); 31212(40); 31232(38) XUMBER 29 169 Proctor, G., & W. Mullings, 22001(38); 22049(46d) Proctor, G., & W. Stearn, 11451(36); 11859(27a); 11923(27a) Purdie, W., 23(46a); 221(46d); 635(38); sn(9); sn(20b); sn(27a); sn(38); sn(40); sn(46a); sn(46b); sn(46d) Quesnay, M. du, 257(27a) Questel, A., 5111(18b); sn(l8b) Ramage, G., sn(l8b) Rehder, A., sn(36); sn(40) Richard, L., 48(27a); sn(l8b) Ridley, H., sn(27a) Riedle, A., sn(43) Robbins, R., 1513(27a); 1514(40); 1516(27a); 1517(27a) Robertson, E., 113(46d); 2996(1a); 3220(40); 5374(27b) Rohr, J. von, sn(l8b) Roig, J., 5054(6a); 14224(27a) Roig, J., J. Acuna & Naranjo, sn(25) Roig, J., & G. Bucher, 53(10); 59(10) Rothrock, J., 87(27a) Ryan, J., sn(l8b) Sagra, R. de la, 647(25); 816(6a); 907(3); sn(25) Sargent, F., 131(30); B131(44c); 613(26); 703(24); 710(43); 3235(24); 3248(30) Schomburgk, R., sn(35) Schubert, B., & H. Winters, 312(43); 401(44c); 406(44c) Schubert, B., H. Winters & I. Velez, 329(43) Senn, H., 331(25) Shafer, J., 187(18b); 3201(22); 3527(44c); 3916(34); 4202(47b); 4394(47b); 7695(25); 7755(17b); 7786(34); 7838(17b); 7984(34); 8012(41); 8135(22); 8171(22); 8420(22); 8436(46b); 8547(32); 8949(44d); 11171(25); 11936(25); 12000(25); 12002(25); 12165 (25); 12254(25) Shafer, J., & Bro. Leon, 13601(25) Shakespear, R., sn(36); sn(40) Shillingford, C., 191(18b) Sintenis, P., 44(43); 327(26); 1343(35); 1484(44c); 2311(43); 2314(43); 2950(43); 3153(43); 5332(44c); 5335(44c); 5782(43); 5832(43); 5827(30); 5829(30); 5845(30); 5910(43); 5935(30); 6096(30); 6568(30); 6590(24); 6755(43); sn(30) Skelding, A., 6705(40); 5450(40); in Adams 10857(27a); sn(40) Skog, L., 1406(43); 1408(24); 1409(33); 1410(27a); 1411(30); 1412(35); 1413(35); 1415(33); 1516(44c); 1517(35); 1539(26); 1586(35); 1542(26); 1544(26); 1583(18b); 1589(33); 1593(44b); 1606(35); 1620(la); 1623(39); 1630(40); 1633(27a); 1634(27a); 1642(27b); 1643(46d); 1731(27b); 1737(41); 1749(33); 1751(16): 1817(18b); 1819(33); 1829(41); sn(33) Skog, L., & G. Proctor, 1636(9); 1639(20c); 1640(12) Skog, L., & J. Skog, 1644(35); 1687a(44c); 1688(43); 1704(26); 1705(43); 1706(43); 1707(30); 1715(30); 1716(24) Skog, L., T. Talpey & D. Pfister, 1608(29); 1610(25); 1615(41); 1616(8) Sloane, H., Horti Sicci 3:27(27a) Smith, H., & G. Smith, 55(18a) Smith, L., A. Hodgdon & F. Gonzales, 3248(25) Spetzman, L., & J. Diaz Colon, 23(43) Squire, F., sn(l8b) Stahl, A., 605b(30); 675(43); sn(43) Stearn, W., 3(la); 4(9); 21(46b); 62(40); lOO(36); 155(27a); 246 (27a); 350(27b); 41 l(40); 442( 12); 446(20c); 469(27a); 491(40); 510(27a); 546(27a); 552(27a); 694(1a); 712(27a); 954(46d) Stehle, H., 565(18b); 568(18b); 2261(18b); 1637(18b) Stehle, H., & M. Stehle, 5173(18b) Stern, W., & D. Wasshausen, 2401(18b) Stevens, F., & 1%'. Hess, 3257(43); 3716(26); 4694(30); 4872(26); 5154(30); 5163(24) Stevenson, J., 547(43) Stevenson, J., & J. Johnston, 2060(30) Stoffers, A., 3172(18b); 3248(18b); 3605(18b); 4255(18b) Stone, M., 117(35); 123(30); 125(46d); 126(20c); 130(24); 132 (33); 134(40); 197(18b); 215(30); 216(27a); 234(27a); 235(29); 350(35); 412(43) Suringar, W., sn(l8b) Swartz, O., sn(1a); sn(l8b); sn(2Ob); sn(27a); sn(37); sn(38); sn(40); sn(41); sn(46d) Talpey, T., lla(30); llb(30); 12(35); 13(37); 14(46d); 15(27a); 16(12); 17(20c); 18(27b); 19(9); 20(46b); 22(40); 23(44~); 25(35); 26(43); 27(30); 28(24); 30(26); 33(33); 34(44b); 35(35); 41(24); 43(29); 62(43); 63(30); 64(30); iI(41); 77(35); 81(16); 92(35); 94(41); 5120(33) Talpey, T., & J. Jimeze, 5120(33) Thompson, W., 7986(40) Turckheim, H . von, 5381(18b) Tussac, F. de, sn(40) Underwood, L., & F. Earle, 235(34); 622(35); 690(35); 708(47b); Underwood, L., & R. Griggs, 803(30) 1062(35); 1174(47b) Valenzuela, J., sn(25) Valeur, E., 569(35) Van Hermann, H., 42(25); 332(25); 1848(25) Van Houtte, Hort. sn(6a) Vtlez, I., 1606(24); 3673(43) Vivaldi, J., 72-8(44c) Wagner, R., 353(35); 412(43); 464(44c); 640(43); 1233(24); 1234 (43); 1664(43) Wall, E., sn(27a) Weaver, R., 1926(46d) Webster, G., 4079(3); 5570(42); 13627(38) Webster, G., J . Ellis & K. Miller, 8087(27a); 8416(9): 8421(27b); Webster, G., & G. Proctor, 5317(40) Webster, G., & K. Wilson, 5017(27a) Wight, A., 130(27a) Wilbur, R., E. Dunn, H. Hespenheide & D. Wiseman, 7871 Wiles, J., sn(27a); sn(40) Wilson, K., & W. Murray, 594(27a) Wilson, N., 463(37): sn(39); sn(4D); sn(46a) Wilson, P., & Bro. Leon, 2847(25); 11617(25) Wolle, J., sn(27a) 94 1 1 ( 1 8b); 9455( 18b) (18b) 170 SMITHSONIAN CONTRIBUTIOSS TO BOTANY ik?rbna, F., sn(l8b) ?ivright, C., 377(32); 382(6a); 647(25); 354(44d); 355(29); 355a (25) ; 1335(32); 3072(19); 3073(3); 3074(17b); 3075(7); 3076 (6a); 3077(25); 3078(45); 3079(4$a); 3080(32): 3080(34); 3081 Yunckei, T., 17404(40); 17781(27a); 18215(2ia), 18349(1a); (35); 3643(17a); sn(17a); sn(22); sn(35) IVjdler, H., 453(30) It?ynter, L., 5669(27a) 18541 (37) \\?right, I\?., sii(27b); sn(38) Wullschlagal, H., 992(46d); 957(27b); sn(27b) TVithout collector, sn(40) Literature Cited Adams, C. D. 1972. Flowering Plants of Jamaica. 848 pages. Mona, Jamaica: University of the West Indies. Allen, G. M. 1939. Bats. x + 368 pages. Cambridge, Massachusetts: Harvard University Press. Ashton, P. S. 1969. Speciation among Tropical Forest Trees: Some Deductions in the Light of Recent Evidence. Bio- logical Journal of the Linnean Society, 1:155-196. GesnPriacees. 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X Achimenantha naegelioides, 161 Achimenes, 158 erecta, 158, 162, 163 longiflora, 161, 163 Amasonia hirta, 162 Anodiscus xanthophyllus, 164 Anthracothorax dominicus, 26-28, 30, 86 mango, 26, 30, 41, 148 viridis, 26, 30 .4pis mellifera, 27 Arbol de navidad, 135 .4rchilochus colubris, 26, 30 Bignoniaceae, 30 Besleria lutea, 159 Bombax malabaricum, 30 Brachyphylla, 30 cavernarum, 27, 31, 32, 135 nana, 27, 32, 144 pumila, 27, 32, 122, 128, 132, 134 Calyptae helenae, 26, 30, 88 Canelon, 52 Car)ophyllaceae, 33 Chiroptera, 29 Chlorostilbon maugaeus, 26, 30, 93 ricordii, 26, 30, 88 swainsonii, 26, 30, 86 hispida, 4, 102 pumila, 4, 44, 152 tenera, 4, 148, 152 Chori$anthera, 4, 5, 44, 146 Clavellina, 95 Codonanthe, 18 Codonophora, 3 grandiflora, 3 lanceolata, 3 albiflora, 134 calycosa, 44, 46, 129 catalpiflora, 128 craniolaria, 131 lessertiana, 128 parviflora, 128 Codonoraphia, 4, 5, 44, 46 Coerebidae, 28 Coleoptera, 27 Columnea, 17 Conradia, 3-5, 36, 43, 46, 98 aureonitens, 162 section Chorisanthera, 3, 4, 6, 145 section Dittanthera, 3, 4, 121 Index (Sew names and new combinations in italics) section Synanthera, 3, 6, 90 calycina, 123 calycosa, 125-127 cassioides, 158 celsioides, 94, 95 clandestina, 124, 127, 128 corrugata, 113, 115 craniolaria, 131 cuneifolia, 4, 109 depressa, 155, 156 exserta, 128 floribunda, 113 fuchsioides, 158 gloxinioides, 146, 147 hispida, 102, 104 humilis, 46, 90, 94, 98 lecontei, 158 libanensis, 102, 113, 115, 118, 119 mimuloides, 148-150 neglecta, 4, 150, 151 pedunculosa, 3, 134-137 var. fI lutea, 134 var. a pallida, 134 pumila, 4, 94, 152 reticulata, 109, 119, 120 fI obovata, 119, 120 scabra, 83 sloanei, 4, 102, 104 verrucosa, 54, 57 ventricosa, $6 fI hngustior, 76, $7 Craniolaria fruticosa, 2, 131, 132 Cpnophaia bicolor, 26 Cyrtandroideae, 1 Digitalis folio oblong0 serrato . , ., 2 Diptera, 27 Drepanididae, 28 Duchartrea, 4, 5, 43 sintenisii, 142 viridiflora, 43, 143-145 Erophylla bombifrons bombifrons, 27, 32, 135 bombifrons santacristobalensis, 27, 32, 141 sezekorni sezekorni, 27, 32, 157 sezekorni syops, 27, 32, 149 Eugesnereae, 3, 36 Enlampis jugularis, 26 Flor de cueva, 110 176 NUMBER 29 177 Gesneria [Gesnera], 1-7, 9-11, 13, 18, 19, 22, 24-32, 34-36, 38-40, 43-46, 51, 52, 54, 56, 57, 94, 97, 98, 112, 120, 127, 134, 146, 147, 158 section Chorisanthera, 45, 100, 145, 147, 151 section Codonoraphia, 121 section Conradia, 6, 46, 90 section Dittanthera, 18, 45, 87, 121, 122, 125, 136 section Duchartrea, 18, 39, 45, 87, 94, 96, 137, 138, 141 section Gesneria, 18, 45, 90, 91, 93, 94, 96, 100 section Lachnoblaste, 45, 85, 87, 122 section Myrrnekianthe, 45, 86, 89, 122, 138 section Pentarhaphia, 6, 18, 29, 45, 46, 48, 51, 77, 78, 80, section Physcophyllon, 29, 45, 90, 93, 100, 105, 111, 115, 145 section Stenochonanthe, 45, 46, 80, 81 acaulis, 2, 4, 8, 11, 13, 15, 16, 24, 30-33, 35: 37, 54, 100-105, 85, 87, 90, 122 107, 109, 113, 119, 158 var. acaulis, 20, 21, 23, 35, 37, 102, 103, 105 var. glabrata, 19, 20, 22, 23, 29, 34, 35, 37, 101, 103, 105, var. grandifolia, 102, 104 107 acuminata, 94, 97, 98 adenantha, 158 affinis, 158 aggregata, 158 albiflora, 134, 137 allagophylla, 158 a lpha , 7, 23, 122, 123, 125, 128 amabilis, 158 amplo digilatis folio tomentoso, 2 angustior, 76 arborescens, amplo flore fimbriato & maculoso, 2, 132 aspera, 45, 46, 48, 49, 59, 67 atrosanguinea, 158 aurantiaca, 158 auriculata, 158 barahonensis, 93, 101, 106, 107, 116, 119 barbata, 15s berteroana, 158 betonicifolia, 158 bibracteata, 158 biflora, 134, 136 binghamii, 47, 49-51, 80 blassii, 158 bonaoana, 54-56 brachysepala, 45, 90-93 bracteosa, 87, 89 breviflora, 158 brevifolia, 46, 50-52 bulbosa, 1% bullata, 159 calcarata, 159 calcicola, 54-56 calycina, 3, 22, 23, 123, 125, 126, 159 calycosa, 7, 10, 16-18, 21-23, 30-32, 35, 37, 123-127 canescens, 159 caracasana, 159 cardinalis, 159 var. a lpha , 122-124 catalpiflora, 128 celsioides, 94, 97 chelonioides, 159 chilensis, 159 christii, 11, 13, 19, 23, 24, 28, 33, 37, 93, 101, 107-109, 113, 116, 117 X chromatella, 159 cinnabarina, 159 citrina, 7, 13, 19, 22, 30, 35, 37, 38, 56, 91, 92, 94, loo clandestina, 12, 18, 21, 32, 37, 123-125, 127, 128 clarensis, 47, 51-53, 62, 67 var. turquinensis, 61, 62 clausseniana, I59 coccinea, 159 cochlearis, 159 confertifolia, 159 cooperi, I59 cooperiana, 159 cordata, 159 coruscans, 159 corymbosa, 25, 40, 42, 43, 128, 159 craniolaria, 3, 131 crenulata, 159 cubensis, 30, 43, 47, 51-57, 61, 65, 67, 69 var. cubensis, 48, 54-56 var. truncata, 17, 48, 53, 54, 57 cumanensis, 159 cuneifolia, 4, 7, 16, 19, 21-24, 30, 32, 35-37, 100, 101, 109-112, 121, 137 cv. El Yunque, 121 cv. Quebradillas, 112 var. disjuncta, 119, 121 var. obovata, 119 cymosa, 74, 75 cynocephala, 159 decapleura, 30, 85, 86, 133 dentata, 159 deppeana, 159 depressa, 155 discolor, 159 dolichostyla, 54-57 domingensis, 54, 56 donkelaariana, 159 donklarii, 159 douglasii, 159 duchartreoides, 17, 23, 30, 54, 86-90, 145 earlei, 159 eggersii, 159 X egregia, 159 ekmanii, 159 elatior, 159 eliptica, 159 elliptica, 159 elongata, 159 eriantha, 159 erubescens, 159 exogonia, 134, 136 X exoniensis, 160 exserta, 3, 7, 13, 32, 36, 37, 79, 121-123, 125, 128-131, 136 SMITHSOSIAS COSTRIBLTIOSS T O BOTANY fascialis, 160 faucialis, 160 fawcettii, 81, 82 ferruginea, 70 filisepala, 67, 69 fimbriata, 131 flacourtifolia, 160 favescens, 160 fragilis, 160 fruticulosa, 160 gardneri, 160 gerardiana, 160 geroltiana, 160 gibberosa, 71 glandulosa, 46, 51, 57-59 glaucophylla, 160 gloxinioides, 32, 39, 145-147 gollmeriana, 160 gracilis, 160 grandis, 3, 160 guazumifolia, 160 guianensis, 160 hahnii, 1FO haitiensis, 12, 37, 45, 47-49, 59, 60, 67 harrisii, 11, 12, 37, 47, 60-62, 65 herbertiana, 160 herminieri, 76 heterochroa, 23, 47, 51, 58, 61-63 heteroclada, 54-56 hirsuta, 160 hispida, 102 hondensis, 113, 160 hookeri, 160 houttei, 160 himilis, 2, 3, 7, 8, 10-12, 16, 19-24, 30, 33, 35, 3i , 44-46, 90, 91, 93-98, 100 humilis, flore flavescente, 2, 95 h>bocarpa, 93, 101, 112, 113 hypoclada, 47, 48, 63, 64 ignorata, 160 inaequalis, 160 incisa, 94, 97, 98 incurva, 50, 160 jamaicensis, 37, 47, 61-65 lanceolata, 47, 48, 65, 66 lasiantha, 160 lateritia, 160 latifolia, 160 Iehmannii, 160 leiocarpa, 150-132 leopoldii, 160 Icptopus, 160 lessertiana, 128 leucomalla, 160 libanensis, 4, 13, 22, 33, 43, 57, 101, 113-115, 118, 119 lindeniana, 160 var. corrugata, 113 lindlepana, $6 lindleyi, 160 lindmanii, 157 linkiana, 160 lobulata, 161 lomensis, 161 longiflora, 76, 161 longifolia, 161 longipes, 161 hlgirostris, 161 lopezii, 113, 114 macrantha, 161 macrorrhiza, 161 macrostachpa, 161 maculata, 161 niagnifica, 161 marchii, 161 melittifolia, 161 merckii, 161 mimuloides, 149 mollis, 161 inontana, 76 montelidensis, 161 mornincola, 54, 53, 69 mortonii, 25, 43, 161 X naegelioides, 161 neglecta, 150 X nigrescens, 161 nigrina, 161 nipensis, 87, 89 nitida, 161 norlindii, 87, 89 oblongata, 161 odontophylla, 47, 48, 52, 65-67 onacaensis, 39, 76, 161 o?7ychocnlyx, 17, 123, 125, 128, 129, 133, 134 organa, 161 oxyphplla, 161 pachpclada, 87, 89 palustris, 161 paluviensis, 161 pardina, 161 parviflora, 128 parvifolia, 46, 48, 49, 67, 68 pauciflora, 7, 19, 21, 25, 30, 32, 37, 91, 94, 98-100 pedicellaris, 8, 11, 13, 14, 16, 19, 20, 22, 28-30, 33, 35, 37, 93, pedunculosa, 3, 8, 11, 16-19, 21, 22, 30, 32, 35-37, 56, 123, pendulina, 162 petiolaris, 162 picta, 162 pilosa, 162 plumeriana, 162 polyantha, 162 portoricensis, 109 prasinata, 3, 162 proctorii, 152, 153 pulchella, 162 101, 115-117 131, 134-137 NUMBER 29 179 pulverulenta, 13, 19, 21, 30, 35, 37, 45, 47, 48, 67-69 pumila, 3, 4, 31-33, 44, 145-149, 152-154, 157 subsp. mimuloides, 16, 32, 148-151 subsp. neglecta, 4, 6 , 8, 22, 32, 37, 148-152 subsp. proctorii, 148, 149, 151-153 subsp. pumila, 21, 37, 148, 149, 151-154, 157 punctata, 162 purpurascens, 33, 58, 161, 112, 115, 117-119 purpurea, 162 X pyramidalis, 162 quadriflora, 128 quadrifolia, 162 quaterniflora, 162 quisqueyana, 140, 141 reflexa, 162 X refulgens, 162 regalis, 162 regeliana, 162 regina, 162 reticulata, 6-8, 13, 28, 32, 33, 35, 37, 49, 93, 101, 106, 111, rhynchocarpa, 162 robusta, 162 rubricaulis, 162 rugata, 162 rupestris, 162 rupicola, 162 rupincola, 162 rutila, 162 salicifolia, 47, 58, 59, 69-71, 73 var. ferruginea, 47, 51, 70-72 var. salicifolia, 51, 70, 71 var. spathulata, 51, 70-72 115, 119-121, 137 salviifolia, 162 samuelssonii, 155, I57 sartorii, 162 saxatilis, 67, 69 scabra, 3, 80-83, 162 var. fawcetti i , 24, 30, 81, 82 var. scabra, 30, 81-83 var. sphaerocarpa, 15, 19, 20, 22, 29, 34, 35, 37, 65, 81-84 var. viridicalyx, 81, 82, 84, 85 sceptroides, 162 sceptrum, 162 schiedeana, 162 schomburgkiana, 162 scopulorum, 54, 55 seemannii, 162 selloi, 163 sellovii, 163 shaferi, 32, 145, 155-157 subsp. depressa, 11, 33, 147, 155-157 subsp. shaferi, 147, 155-157 sintenisii, 140, 142, 143 sphaerocarpa, 83, 84 spicata, 163 splendens, 163 splendidissima, 163 stachydifolia, 163 stachyfolia, 163 stricta, 163 subalata, 54, 55 sulcata, 163 suttoni, 163 sylvatica, 163 sylvicola, 140, 141 tenella, 163 tenera, 152 tetraphylla, 163 tomentosa, 2, 3, 36, 45, 46, 163 trianaei, 163 tribracteata, 163 triflora, 79, 163 trifoliata, 163 truncata, 53, 57 tuberosa, 163 tubiflora, 163 tweediana, 163 ulmifolia, 163 umbellata, 163 uniflora, 163 vargasii, 163 vauthieri, 163 velutina, 163 ventricosa, 3, 4, 7, 12, 16, 24, 25, 30, 43, 45-47, 72-76, 79, 98, 137, 163 f . montana, 76 f. obovata, 76 f. ovata, 76, 77, 79 subsp. cymosa, 74, 75, 78 subsp. ventricosa, 19, 20, 23, 35, 37, 74, 76-79 var. y obovata, 76 var. fi ovata, 76 verdi, 164 verrucosa, 54, 57 verticillata, 164 vestita, 164 viridiflora, 30, 90, 137-139, 143, 144 subsp. acrochordonanthe, 138-141 subsp. quisqueyana, 7, 14, 16, 18, 19, 22, 32, 35, 37, subsp. sintenisii, 19. 20. 32, 35-37, 94, 137-139, 142, 143 138-141 . . . subsp. viridiflora, 21, 22, var. acutifolia, 144, 145 var. colorata, 144 var. obovata, 144, 145 warmingii, 164 warszewiczii, 164 wrightii, 47, 51, 73, 79, 80 xanthophylla, 164 yumuriensis [yamuriensis], zebrina, 164 i2, 62, 87, 96, 138, 139, 144 17-119 Gesneriaceae, 1, 5, 7, 9, 18, 22, 24, 28, 29, 32-36 Gesnerieae [Gesnereae], 1-6, 19, 22, 24-26, 29, 33, 34, 36, 3840, 143 subtribe Conradieae, 3, 36 subtribe Pentarhaphieae, 36 subtribe Rhytidophylleae, 36 180 SMITHSONIAN CONTRIBUTIOKS T O BOTANY Gesnerioideae, 1, 6, 7 Glaucis hirsuta, 26, 30, 74 Glossophaga, 30 Glossophaginae, 29 X Gloxinera, 158 Gloxinia, 158 soricina antillarum, 27, 31, 32, 130, 149 donkelaariana, 159 calycina, 123 lindleyi, 123 sylvatica, 158, 161-163 X Heppiantha naegelioides, I59 Heppiella, 6, 9, 25, 38, 40 corymbosa, 40, 42 cubensis, 40, 43 ulmifolia, 163 verticillata, 164 floribunda, 43, 113 Herincquia [Henrincquia], 4, 5 , 43 Hymenoptera, 27 Kigelia, 30 Kohleria, 158 eriantha, 159 guazumifolia, 160 hirsuta, 160 hondensis, 160 ignorata, 160, 161 lanata, 161 longiflora, 161 longifolia, 160, 161 longipes, 161 mollis, 161 rubricaulis, 162 schiedeana, 162 seemannii, 162 spicata, 163 trianae, 159, 161, 163 tubiflora, 162, 163 ventricosa, 163 vestita, 164 warszewiczii, 162 var. petiolaris, 162 Lepidoptera, 27 Leptonycteris nivalis, 30 Macranthera flammea, 158 Marchantia, 33, 100 Martynia fruticosa, 131 hfelasma scabrum, 161 Mellisuga minima, 26, 30, 41, 42, 81, 148 Mitella, 33 Meliphagidae, 28 Microchiroptera, 29 Mitraria coccinea, 159 Xfonophyllus cubanus ferreus, 141 redmani, 31 redmani clinedaphus, 27, 32, 157 redmani portoricensis, 27, 32, 135 redmani redmani, 27, 32, 149, 151 triflora, 163 hloussonia deppeana, 159-163 Sauti1ocal)x melittifolius, 159 Sectariniidae, 28 Sidulariales, 33 Ophianthe, 4, 5 , 44 Orthorhpnchus cristatus, 26, 30, 93 libanensis, 4, 44, 113 Paliavana, 3 Pentarhaphia [Pentaraphia], 3-5, 7, 36, 43, 46 prasinata, 162-164 section Codonoraphia, 5 , 121 section Duchartrea, 5 , 137 section Eupentarhaphia, 5 , 6, 4G section Synanthera, 5 section l'aupellia, 5 , 121 acaulis, 102 albiflora, 134, 136 calycina, 123 calycosa, 125 catalpiflora, 128, 131 celsioides, 94 clandestina, 127 corrugata, 113 korpmbosa, 42, 164 craniolaria, 131 cubensis, 54, 55 cumanensis, 164 cuneifolia, 109 depressa, 155 tluchartreoides, 87-89 exserta, 76, 128 ferruginea, 70, 72 floribunda, 113 florida, 76, 77, 79 glandulosa, 57, 58 gloxinioides, 146 herminieri, 76, 77 hispida, 102 humilis, 94 incurva, 49, 50 lessertiana, 128, 131 libanensis, 102, 113 lindleyana, 76, 78 longiflora, 3 , 7 , 43, 46, 74, 76, 134, 137 mimuloitles, 148 montana, 76, 77 neglecta, 150 parviflora, 128, 131 pedunculosa, 134 pumila, 152 var. (3 corollae limbo regulari, 54 var. lindleyana, 76 NUMBER 29 181 reticulata, 119 fi obovata, 119 salicifolia, 71, 73 scabra, 83 sloanei, 102 swartzii, 76 tenera, 152 triflora, 73, 79, 80 ventricosa, 76 verrucosa, 54, 55 viridiflora, 144 Pentarhaphieae, 5 Pentarrhaphis, 43 Pheidonocarpa, 6, 7 , 10, 11, 22, 24, 25, 36, 38-40 corymbosa, 7-9, 13, 22, 35, 37, 40, 41, 43, 159, 164 subsp. corymbosa, 21, 41, 42 subsp. cubensis, 41, 43, 161 Phyllonycterinae, 29 Phyllonycteris aphylla, 27, 32, 122, 127, 128, 134 poeyi, 27, 32, 144 Phyllostomidae, 29, 31, 32 Quercus, 131 Rapunculo affinis anomala vasculifera . . ., 2, 103, 104 Rechsteineria, 17, 158 atrosanguinea, 158 aurantiaca, 158 blassii, 158 bulbosa, 158, 159 caracasana, 159 claussenii, 159 cochlearis, 159 confertifolia, 159 coruscans, 159 dentata, 159 discolor, 159, 162 douglasii, 159 elatior, 159 elliptica, 159 erubescens, 159 faucialis, 160 flavescens, 160 fruticulosa, 160 gollmeriana, 160 gracilis, 160, 162 hookeri, 160 lateritia, 160, 161, 163 leopoldii, 160, 163 macrostachya, 160, 161 marchii, 161 merckii, 161 pendulina, 162 polyantha, 162 reflexa, 162 rupicola, 162 sceptroides, 162 schomburgkiana, 160, 162 sellovii, 163 splendens, 163 suIcata, 163 suttonii, 163 trifoliata, 163 umbellata, 163 vargasii, 163 vauthieri, 163 warmingii, 164 subtribq Eurhytidophylleae, 4 Rhytidophylleae, 4, 36 Rhytidophyllum [Rytidophyllum], 3-7, 9-11, 13-15, 18, 19, 22, 24, 25, 34-36, 3840, 45, 46, 76, 138 acunae, 11 auriculatum, 9, 10, 13, 14, 17, 19, 27, 28, 33, 35, 37, 38, berteroanum, 3, 11, 13, 16, 27, 28, 35, 37, 46, 158, 162 bicolor, 39 crenulatum, 159, 163 cumanense, 159, 164 earlei, 25, 159 floribundum, 4, 113 grande, 7, 8, 23, 30, 35, 37, 39, 138, 160 lanatum, 11 leucornallon, 8, 11, 13, 19, 35, 37, 38, 160 lomense, 25, 161 onacaense, 15, 161 plumerianum, 7 rupincola, 25, I62 tomentosum, 2, 7-11, 13, 15, 20-24, 29, 31, 33-38, 104, 105, 158, 160 163 Rousettus aegyptiacus, 30 Sagina, 33 Saxifragaceae, 33 Scrophulariales, I , 9 Sericotes holosericeus, 26, 30, 100 Seymeria cassioides, 158 Sinningia, 17, 158 aggregata, 56, 158, 163, 164 allagophylla, 158, 160-163 barbata, 158 calycina, 123 canescens, 159 cardinalis, 159-161 claybergiana, 160 cooperi, 158, 159 incarnata, 158-160, 162, 163 macrorrhiza, 161 magnifica, 158, 160, 161 regina, 162 sceptrum, 160-162 stricta, 163 tuberosa, 158, 162, 163 tubiflora, 163 verticillata, 160-162, 164 warszewiczii, 164 Smithiantha cinnabarina, 159 geroltiana, 160 X hybrida cv. Sulphurea, 159 182 multiflora, 158 zebrina, 162-164 cv. Exoniensis, 160 cv. Refulgens, 162 cv. Robusta, 162 Spanish Tea, 135 Trochilidae, 28, 30 Trochilus polytmus, 26, 30, 41, 81, 148 SMITHSONIAS COSTRIRUTIOSS TO BOTAKY Tupa acuminata, 134 Vanhouttea gardneri, 160, 161 salviifolia, 162 Vaupellia, 4, 44, 125 calycina, 4, 5 , 44, 123 Yerba parrera, 110 * U.S. GOVERNMENT PRINTING OFFICE: 1975-586-384/43