’ ‘ ”3""- {of '-n- '{t‘ ,‘J '7 ’55"? I '(fiir‘f “-1 Mon-.1 :W& ’r‘l -Ts‘ “14,4 «1%.:{t‘flku} 4. 2‘ x “w‘ ”‘1‘ \. . C ”"5 L‘V‘IA‘. ,~ ‘3; {fl “3‘ . “3.. ‘- -~ ‘ x mix. ”-22 . 75' < .4: av .r'v'. .Jr' 0'“. - I. -WDV \( 'Wfixx‘. ‘ «v. rope... ‘Uom-Ld‘idky)~ min.» - 31'3": '4"! 1.- In"; ):,}n'n {,12 I" 141% .K V is: . I. (’7 ; > '4‘)! .l I - yum“ ...r!I-"" % 9w". sch”)? 2'- 7': )5‘9' tux“: f” I: 1:3,};1' ’ll 7‘ 7:4’ " -"'/- a :TXN.) -'C"' flame. _< '1' ‘A .- .2 ('K, 4' .,I 2 . . . . 4 1/", . _.I {(55954 4‘“.j{ . '. I, r 4 _ ’y . .I 2‘ :‘.‘!.';’r"' I," ‘ ‘. h (.l ,: 'Ji'f/ '3” {fl , ’ Hf: 1.,427. . .Ir‘ .. .. 2_ ,_ . _ 1"," x" .1: “,."' M: Jr“ . '-:';. ' f“ , ‘ l ' 2 m , ,g . ”.513; . {4'32 tr. 73w}; 1 V .. a -' 'v' ' I", ' ' ‘ , I}: 2416‘” ~ I: I 2 "r“: ' v." ."',, ' ’« - 1"}; #3: ’1': ’34 . I r 1"" '1; r. x r- 1.7;:- 19” 7 , 22$ng 1‘ “I" ( ”uh I . Am 7' 2;! "92:1" ' .r' ':.',;.'i-' , } ' H" "1?? v' ’3" 3‘“ ’3" “('1‘ ." ,1“; ; ; :- 57.2, '1 “ht," ~l, ‘ '7'! ‘1 ‘g/l . V'GQI" V: ‘ 0‘ l V . C . I . ' v rt '7 v 2 . . 5" us, - . 1:} .393" '36” 5'! 2 ‘ . .‘ ' - y w»; 'A - ‘ w"; 1. ”fife-""3"" IM': -/ l 2 . . x _ . , 2 a: _ :33!erch 3" “7.1; ’j"' .. jar? 6,1“:ngflutfjxm 1 ‘ ' ' . ' ' 4 .v/V. ’ 34,-" m. ’ I, 1‘" '-v' 35")“: 1' 31”.: . Ell-1'"; , 9 . UL'M‘} "'39:"? fit-”y'a- v y” .4 ’ ‘ ' . 'l .' ,‘ '2 51'7"" if 'r’lj', "9’1‘ 7 I; 5"?“ ’ :54!" 1- .. ' s, J?! :‘JX J pf". . , $23.0" ' "I ‘.' .53; all]? {nfigfou’if' W .‘."$.I;,7'5' 7.5:: 'II' "'J' o-a u "5%?31. ' Wm H ’ . 37.3: ' m: army, NI. 'l "'1... Ritz r'" I I, 234/51 (17.31 g.“ I; [/1 LIBRARY Michigan State University This is to certify that the thesis entitled Revision of Medinilla (Melastomataceae) of Borneo presented by Jacinto C. Regalado, Jr. has been accepted towards fulfillment of the requirements for Master OLSLiemdegree in Botany /,6Z‘fl/W Major professor Date ¢d Md? [yfgl / / 0-7639 MS U is an Affirmative Action/Equal Opportunity Institution i MSU LIBRARIES - RETURNING MATERIALS: Place in book drop to remove this checkout from your record. FINES will be charged if book is returned after the date stamped below. REVISION OF MEDINILLA (MELASTOMATACEAE) OF BORNEO By Jacinto C. Regalado, Jr. A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Botany and Plant Pathology 1988 ABSTRACT REVISION OF MEDINILLA (MELASTOMATACEAE) OF BORNEO BY Jacinto C. Regalado, Jr. Fifty-two species of Medinilla are now known from Borneo, 30 of which are described as new. Twenty-two taxa are known only from one to three collections. Twelve species groups have been recognized and defined. A more thorough understanding of the genus awaits further study of Philippine and New Guinea materials. A key to the Bornean species, illustrations, distribution maps, and ecological notes are provided. Four previously described species are recorded for the first time for Borneo: Medinilla succulenta (Blume) Blume, fl. pterocaula Blume, fl. varingiifolia (Blume) Nayar, and M. Qhwii Nayar. One new combination and four reductions have been made. Medinilla tawaensis Merrill is transferred to Catanthera; fl. caudatifolia Schwartz and M- hasseltii Blume var. subsessilis Schwartz are reduced to m. crassifolia (Reinw. g5 Blume) Blume; u. dajakorum Schwartz and M- motleyi Hook. f. g; Triana are conspecific with M- gorallina Cogn. and M- macrophylla Cogn., respectively. ACKNOWLEDGMENTS I am deeply grateful to my major professor, John H. Beaman, for the inspiration and guidance through the course of my studies at Michigan State University. I thank Peter Murphy, Frank Ewers, and Michael Price for counsel and for reviewing the manuscript and Wendy Bromley for the excellent illustrations. Her work was supported by the William T. Gillis Memorial Fund. The Latin diagnoses were edited by Dr. Mladen Kabalin. I am indebted to the directors and curators of herbaria that loaned specimens and provided space for study. Arnold Arboretum, Harvard University, Cambridge (A), Botanische Garten und Botanisches Museum Berlin-Dahlem (B), Bernice P. Bishop Museum, Hawaii (BISH), British Museum (Natural History) (BM), Herbarium Bogoriense, Bogor (BO), Jardin Botanique National de Belgique, Meise (BR), Institut fur Allgemeine Botanik und Botanischer Garten, Hamburg (HBG), Royal Botanic Gardens, Kew (K), Department of Botany, University of Malaya, Kuala Lumpur (KLU), Department of Botany, Faculty of Science, Kyoto University (KYO), Rijksherbarium, Leiden (L), University of Michigan Herbarium, Ann Arbor (MICH), Missouri Botanical Garden, St. Louis (MO), Beal-Darlington Herbarium, Michigan State University, East Lansing (MSG), New York Botanical Garden, New York (NY), Philippine National Herbarium, Manila (PNH), Forest Department Herbarium, Sandakan (SAN), Botanic Gardens, Singapore (SING), University of California, Berkeley (UC), Institute of Systematic Botany, Utrecht (U), Smithsonian ii Institution, Washington, D.C. (US). Herbarium symbols are used in the citation of specimens according to Holmgren §£_al. (1981). The study was supported by National Science Foundation grants BSR-8311352 and BSR-8507843, John H. Beaman, Principal Investigator. Additional funding was provided to me by an American Society of Plant Taxonomists Travel Award. iii TABLE OF CONTENTS LIST OF TABLES ............................................ LIST OF FIGURES ........................................... MORPHOLOGY ................................................ Vegetative Morphology ................................ Floral Morphology ..................................... Fruit and Seed Morphology ............................. SYSTEMATIC TREATMENT ...................................... Synonymy, distribution and generic description ........ Species alliances ..................................... Key to species ........................................ 1 2 U . Medinilla speciosa (Reinw. gg Blume) Blume ......... . Medinilla guadrifolia (Blume) Blume ................ . Medinilla kinabaluensis Regalado ................... . Medinilla urgphylla Stapf .......................... iv 10 ll 13 13 15 16 l6 16 17 23 3O 31 34 36 11. 12. 13. 14. 15. 16. 17. 18. I9. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. . Medinilla rubrifrong Regalado ...................... . Medinilla pterocaula Blume ......................... . Medinilla atroviridis Regalado ..................... . Medinilla clemensiana Regalado ..................... . Medinilla corneri Regalado ......................... 10. Medinilla danumensis Regalado ...................... Medinilla beamanii Regalado ........................ Medinilla allantocalyg Regalado .................... Medinilla fragilis Regalado ....................... Medinilla latericia Regalado ....................... Medinilla pedunculosa Regalado ..................... Medinilla crassifolia (Reinw. ex Blume) Blume ...... Medinilla laxiflora Ridley ......................... Medinilla botryocarpa Regalado ..................... Medinilla longipedunculata Cogn. ................... Medinilla sessiliflora Regalado .................... Medinilla richardsii Regalado ...................... Medinilla montisaping Regalado ..................... Medinilla muricata Regalado ........................ ugg13111§_hgmgg§ggrg Regalado ...................... Medinilla rayae Regalado ........................... Medinilla subauriculata Regalado ................... Medinilla salicina Ohwi g; Regalado ................ Medinilla succulenta (Blume) Blume ................. Medinilla guadrialata Regalado ..................... V 37 38 39 41 42 43 46 48 50 51 52 54 55 56 S9 60 63 64 65 66 69 71 74 76 77 30. Medinilla myrmecorhiza Regalado .................... 79 31. Medinilla aggregata Bakh. f. ....................... 80 32. Medinilla suberosa Regalado ........................ 81 33. Medinilla amplectens Regalado ...................... 83 34. Medinilla varingiifolia (Blume) Nayar .............. 85 35. Medinilla ohwii Nayar .............................. 88 36. Medinilla macrophylla Blume ........................ 88 37. Medinilla rufescens Regalado ....................... 91 38. Medinilla corallina Cogn. .......................... 91 39. Medinilla lasioclados Stapf ........................ 93 40. Medinilla endertii Regalado ........................ 94 41. Medinilla rufopilosa Ohwi gg Regalado .............. 95 42. Medinilla stephanostegig Stapf ..................... 97 43. Medinilla alternifolia Blume ....................... 98 44. Medinilla formanii Regalado ........................ 99 45. Medinilla decurrens Cogn. .......................... 102 46. Medinilla kemulensis Regalado ...................... 103 47. Medinilla serpens Stapf ............................ 106 48. Medinilla capillipes Regalado ...................... 107 49. Medinilla flagellata Stapf ......................... 109 50. Medinilla lorata Stapf ............................. 110 51. Medinilla sandakanensis Regalado ................... 111 52. Medinilla brevipedicellata Regalado ................ 114 Excluded Taxa .......................................... 114 Dubious Species ........................................ 115 vi OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO vii 116 116 119 128 LIST OF TABLES Table Page 1. Morphological characters examined for Medinilla. .......... 12 2. Differential characters of the Medinilla species alliances ................................................. l8 viii LIST OF FIGURES Figure Page 1. Distribution of species of Medinilla section Medinilla ..... 33 2. Medinilla kinabaluensis Regalado ........................... 35 3. Medinilla atroviridis Regalado ............................. 40 4. Distribution of species of the M. corneri alliance ........ 44 5. Medinilla beamanii Regalado ................................ 47 6. Distribution of species of the M. beamanii alliance ........ 49 7. Distribution of species of the M crassifolia alliance ..... 57 8. Distribution of species of the M. sessiliflora alliance .... 62 9. Distribution of species of the M. myrtiformis alliance ..... 67 10. Medinilla rayae Regalado ................................... 70 11. Medinilla subauriculata Regalado ........................... 73 12. Medinilla salicina Ohwi g; Regalado ........................ 75 13. Distribution of species of the M. succulenta alliance ...... 78 14. Medinilla amplectens Regalado .............................. 84 15. Distribution of species of the M. varingiifolia alliance ... 87 16. Distribution of species of the M. macrophylla alliance ..... 90 17. Distribution of species of Medinilla section Heteroblemma . 100 18. Medinilla kemulensis Regalado ............................. 105 19. Medinilla capillipes Regalado ............................. 108 20. Medinilla sandakanensis Regalado .......................... 113 ix INTRODUCTION Medinilla (Melastomataceae) is a genus of epiphytic and terrestrial shrubs and climbers of the paleotropics. It includes about 400 species (Shaw, 1973) distributed in Africa, Madagascar, India, Ceylon, Burma, Indochina, southern China, Thailand, Malay Peninsula and eastward to the islands of the Malay Archipelago, New Guinea, northern Australia, Micronesia, and Melanesia. It is by far the largest of all melastome genera occurring in Malesia, a floristic region made up of the Malay Peninsula and islands of the Malay Archipelago extending to New Guinea. About 200 species are known from this region, which is the center of diversity of the genus. The objective of this study is to document the diversity of Medinilla in Borneo by this first attempt at a revision of the Bornean species. It is hoped that subsequent studies in the field and laboratory will provide better understanding of evolutionary trends and relationships of the species than is possible with the limited material now available. HISTORICAL ACCOUNT Medinilla was established in 1826 by Charles Gaudichaud-Beaupre, a French naturalist and circumnavigator. The type species is M. rosea which was collected during an expedition to the SW Pacific. The genus is named in honor of Jose de Medinilla de Pineda, once governor of the Marianas (Ladrones) Islands. Carl Ludwig Blume, a German-born Dutch botanist and long-time I)irector of the Rijksherbarium, made the first extensive study of the genus. His initial publication on the Melastomataceae appeared in the Bijdragen tot degflora van Nederlandsch Indie (1826), where Medinilla ‘was included in Melastoma. It was not until 1831 when his paper, Egg; leinige ostindische und besonders avanische Melastomaceen, was jpublished in Flora (vol. 14) that he separated Medinilla and nine other ,genera from Melastoma. Blume’s species were classified under 4 sections (Campsoplacuntia [- Medinilla], Sarcoplacuntig, Hypenanthe, and Dactyliota). Blume (1849) subsequently published a two-volume work on the Melastomataceae in which he described 3 new species of Medinilla under section Sarcoplacuntia, elevated sections Hypenanthe and Dactyliota to generic rank, and established 2 new sections, Heteroblemma and Apateon. The genus also has been considered in monographic and floristic works of Naudin (1851), Miquel (1855, 1860), Triana (1871), Cogniaux (1891), and Krasser (1893). Naudin's treatment added little to the knowledge of the genus already provided by Blume. He recognized 12 of Blume's species but was unable to study 17 others, presumably because of the unavailability of specimens. Miquel accepted Naudin's treatment in his Flora van Nederlandsch Indie and described 3 new species, bringing the total to 37 species known at that time in the former Dutch East Indies. Cogniaux wrote the worldwide treatment of the Melastomataceae in De Candolle's Monographiae phanerogamarum in which close to 100 species were enumerated from the Old World. Cogniaux also worked out the family in Boerlage's (1890) Handleiding tot de kenntnis der Floraivan Nederlandsch Indie. This included only generic descriptions but a list of about 41 species known to occur in the former Dutch East Indies was included. In the late 19th century Otto Stapf at Kew became a principal authority on the Melastomataceae. He described three new species of Medinilla from the collections by Haviland on Mount Kinabalu (Stapf, 1894) and three others from Sarawak (Stapf, 1895). Merrill (1929) described one new species from Elmer's collections in the eastern part of the former British North Borneo (Sabah). Schwartz (1931) described two new species from Central Borneo (Dutch Borneo), a vast area that is still relatively unexplored. The Philippine species of Medinilla were extensively studied by Merrill (1913) at the Bureau of Science and by Elmer who worked independently and published in his Leaflets of Philippine Botany. In 1923-26, Merrill published the Enumeration 0 Phil ine Flowerin Plants wherein he recognized over 100 endemic species of Medinilla (Merrill, 1923). Representatives of the genus in New Guinea were studied by E. G. Baker, Jr. (1914) who examined the collections of the Wollaston expedition and by Mansfeld (1927) who worked in northeastern New Guinea (former Kaiser-Wilhemland). Species from the Malay Peninsula were studied by Stapf and King in the Materiala for a Flora of the Malay Peninsula (1900), by Ridley in the Elaga of the Malay Peninsula (1922), and recently by Maxwell (1978). In the 19303, R. C. Bakhuizen van den Brink, Jr. (1943) prepared a monograph of the Melastomataceae in Southeast Asia. Due to prevailing war conditions he based the study only on specimens at Utrecht and Leiden. In the introduction he noted that collections from Dutch Borneo had been distributed to only limited extent by the Buitenzorg (Bogor) herbarium. He also suggested that the genus might be segregated into two or more genera when more Philippine material was studied. During the period when Bakhuizen van den Brink, Jr. was writing his treatment, intensive exploration was undertaken in the rich flora of Papua New Guinea, particularly the Archbold expedition, which resulted in more novelties (Merrill & Perry, 1943; Ohwi, 1943). Ohwi put several in scheda names on herbarium sheets of Bornean specimens but these were never published. The manuscript that Ohwi was working on at the time of his death has not been found (J. F. Veldkamp, personal communication). OVERALL GEOGRAPHIC DISTRIBUTION Medinilla has a bicentric distribution pattern. The two centers, Africa and Asia, have no known species in common and far more species occur in Southeast Asia than in Africa. In Africa the genus is distributed mainly south of the equator along the tropical rain forest belt. Most of the species are found in Madagascar and the east coast of Africa. The distribution pattern shows that the genus is adapted to warm and humid forest environments. The Sahara desert in the north and the Kalahari desert in the south act as barriers. The Asian species are also found in warm and humid forest habitats. In Asia the distribution of the genus is in the Indian-Himalayan ranges southward to Burma, Thailand, Indochina, Malay Peninsula, and eastward to the islands of the Malay Archipelago, New Guinea, Polynesia, and northern Australia. Both Nayar (1972) and Cheih (1983) postulated the origin of the genus in Gondwanaland (South America, Africa, India, Australia, and Antarctica). By the end of the Cretaceous, these land masses had drifted apart. Cheih (1983) suggested that after the Indian plate drifted northward and collided with the Laurasian plate in the early Tertiary, several species of Medinilla successfully dispersed across southern Asia and ultimately migrated eastward to the Malesian islands, the Western Pacific and northern Australia. Populations of many original species that were established along the Himalayan region may have succumbed to the deterioration of climate in the Pleistocene and became extinct, leaving a few isolated disjunct species on the warmer southern flank of the Himalayas. The African species were forced by the desertification of the Sahara to move south of the equator into the surviving patches of rain forest. BORNEAN DISTRIBUTION Borneo is the largest island in the Malay Archipelago and the third largest in the world. In area it approximates 739,175 square kilometers. Borneo is divided politically into Sabah and Sarawak, which belong to the Federation of Malaysia, Brunei, an independent sultanate, and the largest portion, Kalimantan, which is part of Indonesia. The size and equatorial position of the island, the high temperature and humidity, the variation in seasonal rainfall, and the range in altitude are favorable conditions to the development of an exceedingly rich and diversified flora (Merrill, 1930). The island is 6 traversed by long mountain ranges, including the Crocker Range culminating in Mount Kinabalu (4101 m), the highest peak between the Himalayas and New Guinea. Merrill (1921) listed 4,924 species of flowering plants credited to Borneo, representing 1,152 genera and 157 families. Masamune (1942) made a similar compilation and brought the total up to 7,201 species distributed in 1,310 genera and 165 families. On the basis of a conservative estimate by Merrill (1950), the flora of Borneo is somewhere between 12,000 to 15,000 species. Merrill's list included 20 species of Medinilla. There is no botanical exploration documented in Borneo prior to Korthal's pioneering work in 1836. Subsequent botanical reconnaissances were made by Odoardo Beccari, a distinguished Italian botanist, who came to Sarawak in 1865 and John Whitehead, a British ornithologist. Important collections were made between 1851-1900 by Low, Motley, Lobb, Burbidge, Hallier, Haviland, Hose, and Nieuwenhuis. Botanical exploration in the present century has been considerable although still highly inadequate. Collections of great importance have been made by Winkler, Endert, Gibbs, the Clemens, Topping, Elmer, Kostermans, and the foresters of the Sabah and Sarawak Forest Departments as well as botanists involved in the Oxford University and Royal Society Expeditions (van Steenis-Kruseman, 1950). In spite of the efforts of these and other collectors, the flora of a major part of Borneo (Kalimantan in particular) is represented by one of the lowest collection densities of herbarium specimens of any place in the world. In 1972 194,200 herbarium specimens had been collected, the equivalent of only 26 specimens/100 square kilometers (Prance, 1978). Of the 52 Bornean species of Maglallla recognized in this treatment, 41 are presumably endemic. Most of the species are rare, very local, and 22 of them are known from one to three collections. Borneo shares a few of the common species with the Malay Peninsula, Java, and Sumatra, but no species occur in both the Philippines or Sulawesi and Borneo. Borneo is rivaled by the Philippines and New Guinea in terms of diversity of Medinilla.. Merrill (1917) noted that a very high percentage of Philippine Medinillas are not only endemic but also very local. The Philippine situation is often a result of insular isolation, while endemism of Medinilla in Borneo is mostly associated with edaphic and altitudinal factors. Several species of section Meteroblemma, for example, are exclusively confined to limestone hills in Sarawak. Mount Kinabalu, noted for its flora of high specific endemicity, harbors 17 species, 8 of which are endemic. Some species on Mount Kinabalu can be found also in lower elevation but floristically similar mountains such as Alab in Sabah, Mulu in Sarawak, Raya in Central Kalimantan, and Kemul in East Kalimantan. Medinilla ranges from coastal and low elevation riverine forests to mossy and mid-elevation montane forests. However, there is little apparent unity in distribution patterns of related species, perhaps in part because of the inadequacies of collections from Kalimantan. Sabah, Sarawak and Brunei are relatively better collected. No information is available on pollination and seed dispersal for Medlnilla. The flowers are not scented and do not provide nectar, but perhaps the brightly colored flowers and showy bracts in certain species attract pollinators that collect pollen. The fruits that ripen 8 with red and fleshy pericarp are likely eaten and the seeds dispersed by birds. SYSTEMATIC POSITION AND TRIBAL RELATIONSHIPS The Melastomataceae are a large, predominantly tropical family of some 200 genera and 4000 species (Cronquist, 1981). It ranks as the seventh largest family of flowering plants (Wurdack, 1986) and the second largest family in the order Myrtales (Cronquist, 1981). The family is best developed in South America; two-thirds of the known species occur in the New World tropics (Nayar, 1972). Traditionally the family has been divided into 3 subfamilies (Astronioideae, Melastomatoideae, and Memecyloideae) and 14 tribes (Almeda, 1978). No genus occurs both in the New World and Old World. Medinilla belongs to the subfamily Melastomatoideae, tribe Dissochaeteae. At present there are several unresolved problems on tribal relationships (Veldkamp, 1978). Various authors have presented diverse schemes of classification and interpretation of the tribe (Triana, 1871; Cogniaux, 1891; Krasser, 1893; Bakhuizen van den Brink, Jr., 1943; Maxwell, 1980; van Vliet, 1981). The most extensive study of the tribe was made by Bakhuizen van den Brink, Jr. (1943). He pointed out that the usual subdivision of the Dissochaeteae based on the length of the stamens and the characters of the connective were of no value. He also regarded Medinilla to be a heterogeneous group. Backer and Bakhuizen van den Brink, Jr. (1963) suggested that the degree of concrescence between the calyx tube and the ovary and the depth of the extraovarian chambers can help delimit the various genera in the tribe. 9 It is beyond the scope of the present study to address problems in tribal and generic delimitation, but it is important to point out some of the characteristic features of closely related genera, namely Carionia, Catanthera, Hypenantha, Eachycentria, Plethiandra and Pogonanthera. Medinilla differ from the above mentioned genera by having anthers with a dorsal crest or keel or spur on the connective and a pair of basal lobed extensions, referred to as ventral appendages, at the base of each pollen sac. The monotypic genus Carionia of the Philippines closely resembles Medinilla in habit, but it can be distinguished by the long and narrow calyx lobes. Pachycentria differs from Medinilla in the absence of a dorsal spur and extraovarian chambers while Eoggnanthera is well characterized by the presence of tufted hairs at the back of the anthers in place of a dorsal spur. The latter is often confused with Pachycentria, but the non-tuberous roots and biauriculate leaves distinguish it in sterile state. Hypenanthe is a segregate from Medinilla (Blume, 1849) but was not recognized by many authors until the time of Bakhuizen van den Brink, Jr. At least four species of the genus are distinguished by large pilose deciduous bracts and a furfuraceous or pilose calyx tube. In having anomalous xylem Mederella (- Catanthera) provides a close link with section Meteroblemma, but it differs in other characters as noted by Nayar (1966). The infrageneric classification is not yet fully resolved. Bakhuizen van den Brink, Jr. (1943) did not recognize Blume's four sections and created instead two new sections, Eumedinilla and Meteromedinilla. These two sections differ in a) thickness of the calyx wall, b) length and shape of stamens, and c) length of the ovary 10 in relation to that of the calyx tube. This classification, however, was not satisfactory for the Bornean species so I have recognized 12 informal species groups, two of which correspond to sections recognized by Blume. A more formal infrageneric classification must take into account the species from other geographic regions. SPECIES CONCEPT Because the only sources of information in this revision are herbarium specimens and past taxonomic concepts, the species concept is a traditional morphological one. The criterion of reproductive and genetic isolation that defines a biological species concept (Mayr, 1970) cannot be applied as there are no available data on pollination biology, chromosome numbers, population dynamics, hybridization and polyploidy. Mishler and Donoghue (1982) suggest a pluralistic outlook on species and urge systematists to develop species concepts for their particular taxonomic groups. The specimens were sorted geographically into homogeneous and mutually distinct entities, followed by analysis and evaluation of taxa (Leenhouts, 1968). To facilitate recognition of distinct entities, I employed the species-standard method (Rollins, 1952) using previously described species as biological standards of comparison. The taxonomic judgment depended in part on my experience with allied species occuring in the neighboring regions which were compared and related to the Bornean species. Specimens that had the same pattern of definitive characteristics were grouped together. In Medinilla variability is greatest among species that are widespread. These are represented by relatively numerous specimens which cannot be readily distinguished. 11 On the other hand, many species of Medinilla tend to be highly localized or restricted in distribution. In Borneo the species of Medinilla are nearly parallel in their degree of distinctiveness, hence no infraspecific categories were assigned. The flora of Borneo is so poorly known that recognition of geographical subspecies or varieties would be of little taxonomic relevance. MATERIALS AND METHODS This revision of the Bornean species of Medinilla is based upon a study of both herbarium specimens and the literature. Names published under Medinilla were reviewed from the Kew Index, Merrill's (1921) enumeration, and various floristic and monographic works. Specimens were borrowed from herbaria that have significant collections from Borneo. About 1500 specimens representing 690 collection numbers of Medinilla were examined. The specimens were sorted geographically, then by collector. Specimens were systematically examined for the morphological characters listed in Table 1. Analyses were made on boiled flowers and fruits using a Zeiss dissecting microscope fitted with an eyepiece micrometer. Measurements were taken for all specimens that represented a particular taxon when ten or fewer specimens were available. In the case of taxa for which more than ten specimens were available, measurements were taken from specimens that had flowering and/or fruiting materials. Data on the distribution of Medinilla in Borneo were gathered from specimen labels and entered into a database file (MEDINILA) using a microcomputer database management system, dBASE I I Plus. Data include geographic as well as altitudinal distribution, abundance in terms of 12 Table 1. Morphological characters examined for Medinilla. l. Habit a. b. creeper, shrub, treelet epiphytic, terrestrial c. height 900‘“ WHHUOQMCDO-OU‘O) . Stem . shape (cylindric, quadrangular, winged) texture (smooth, pustulate) . pubescence (glabrous, furfuraceous, etc.) diameter . Leaves phyllotaxy (alternate, opposite, whorled) nervation (3, 5, 7, 9 -nerved or -p1inerved) petiole (present, absent) petiole length leaf shape leaf length leaf width leaf apex leaf base leaf margin leaf axils (glabrous, pilose, tufted) Inflorescence MOO-00‘” OQH'lGO-OO‘D a. b. type (solitary, cymose, umbellate, paniculate, glomerulate) position (axillary, terminal, subterminal) number of flowers (few- or many- flowered) bracts and bracteoles . pubescence . peduncle and pedicel . Flower . calyx color, shape, size, indument corolla color, shape, size stamen number, condition (equal or unequal) anther shape and length, appendages . depth of extraovarian chambers style length shape of stigma . Fruit and seed fruit shape, texture, diameter, color seed shape, size, color 13 collection frequency, vernacular names, and economic importance. A program (CITATION) written by the author was used to prepare specimen citations, index to exsiccatae, and determination labels. This program was modified from the LABELS3 collection database software (Regalado et al., 1987). It is also compatible with the Mount Kinabalu database project at MSG, wherein records of specimens collected on this mountain can be extracted and incorporated into the MEDINILA file without rekeying the data. A list of specimen citations is on file in the herbaria at Kew, Leiden, Arnold, and Michigan State University. Dot maps were used to show distribution of taxa. Place names on labels without latitude and longitude information were located on the map using standard gazetteers for Malaysia, Brunei, and Indonesia (U.S. Board on Geographic Names, 1970, 1982). MORPHOLOGY Vegetative Morphology The species of Medinilla in Borneo are epiphytic or climbing shrubs, treelets, or creepers. Medinilla is reported to have 75 percent of total species that are epiphytes (Kress, 1986). A few specimens have been recorded as large trees, but they were probably epiphytes mistaken as trees. Some collectors have noted them as parasites, but parasitism is unknown in other Myrtales (Cronquist, 1981). The stems are either cylindric or quadrangular. Certain species have distinctive winged branches, at least in the juvenile stage. Mature branches are more or less slender since the maximum diameter rarely goes beyond 10 mm in most herbarium specimens. The stem surface may be smooth or pustulate, glabrous or with varying forms and degrees 14 of pubescence. Adventitious roots often grow from leaf axils or defoliated nodes which are sometimes swollen. Species of section Heteroblemma have wood with lobed xylem (van Vliet, 1981). The nature of the leaves provides several features that are most useful in distinguishing the species. Leaves may be alternate (section Heteroblemma), verticillate (section Medinilla) or opposite. Opposite phyllotaxy is the most common condition. The venation of the leaves consists of subparallel longitudinal nerves (primaries) that range in an odd-numbered fashion from 3-11. Leaves are referred to as nerved when all primaries arise from a common point at the base of the blade. Leaves are referred to as plinerved when one or more pairs of inner nerves diverge from the midvein at a point above the leaf base. The midrib and lateral nerves are generally impressed on the adaxial surface and raised on the abaxial surface. Transverse veins run across the blade perpendicular to the midrib. The relative conspicuousness of transverse veins and the degree of reticulation are useful in distinguishing the species. Leaf shape and size range from large and rotund leaves of Medinillaikemulenala to small and lanceolate leaves of M. richardsii. The leaves of Medinilla, in general, are elliptic, coriaceous and essentially glabrous. Pubescence on the leaves is observed only in species of section Heteroblemma and in the M. macrgphylla alliance. The leaves are most commonly entire, except for some species in section Heteroblemma that have small serrulations. The leaf axils are generally glabrous but may be pilose, or tufted with bristles as in M. speciosa, M. stephanostegia, and M. muricata. The absence or presence of a petiole, except in a few cases, is a useful character in distinguishing species. 15 Floral Mor holo The inflorescence is derived from the basic cymose type that may be fascicled or glomerulate, umbellate or paniculate and few- to many- flowered. In some species the flowers are solitary or paired. Inflorescences are often lateral in position, arising from leaf axils or from leafless nodes. Terminal inflorescences are exhibited in M. speciosa, M. stephanostegia, M. ohwii and M. varingiifolia. Only M. speciosa and M. stephanostegia display leafy and showy bracts. Bracteoles subtend the individual flowers and are often subulate and small (1 mm long), and persistent or caducuous. Flowers of Medinilla are ephemeral. Flowering material is scarce and often collected in the advanced stages. The flowers are 4-, 5-, rarely 6-merous. The number of floral parts was found unreliable in differentiating the species. The calyx (hypanthium) varies in shape from campanulate to urceolate and ovoid, is often red in color and is generally glabrous, except in M. serpens, M. capillipes, and in the species of the M. macrophylla alliance. The rim may be truncate or marked with 4 or 5 calyx teeth. The corolla consists of 4 or 5 petals, rarely 6, that are white to pink to red and obovate. The stamens vary in number from 8 to 10 or 12, are equal or unequal in size, and open by a single terminal pore. A connective generally is not produced at the base. The pistil consists of a 4-5 (-6) celled ovary with numerous ovules axially attached to the placenta, a slender terete style, and a punctiform or minutely capitate stigma. 16 Fruit and seed morphology The fruit is technically a berry that is often globose in shape, sometimes cupuliform or cylindric. It is generally glabrous except in Medinilla serpens, M. capillipes and M. brevipedunculata and some species of the M. macrophylla alliance. The pericarp may be thick (section Medinilla) or thin (species allied to M. succulenta). Seeds of Medinilla are generally minute (0.5-1.5 mm long), cochleate to ovoid in shape, yellow to orange in color; the testa may be smooth or reticulate. The seeds have a conspicuous lateral raphe. SYSTEMATIC TREATMENT MEDINILLA Gaudich., Voy. Uranie (1826) 484; DC., Prodr. 3 (1828) 167; Blume, Flora (1831) 464; Mus. Bot. Lugd.-Bat. 7 (1849) 17; Naudin, Ann. Sci. Nat. III, 15 (1851) 285; Triana, Trans. Linn. Soc., London 28 (1871) 85; Cogn., DC. Monogr. phan. 7 (1891) 572; Bakh. f., Rec. Trav. Bot. Neerl. 40 (1943) 147. - Diplogenea Lindley, Quart. J. Sci. Arts 2 (1828) 122. - Triplectrum D. Don a; Wight & Arn., Prodr. (1834) 324. - Erpetina Naudin, Ann. Sci. Nat., III, 15 (1851) 299. - Cephalomedinilla Merrill, Philip. J. Sci. 5 (1910) 204. Distribution. About 400 species in tropical Africa, Madagascar, India, Ceylon, Burma, Indochina, S China and SE Asia throughout Malesia to N Australia and Polynesia. Epiphytic and terrestrial shrubs, erect, scandent, or creeping. Branches generally cylindric, smooth or pustulate. Leaves alternate, 17 opposite, or verticillate, sessile or petiolate; blade fleshy or coriaceous, generally elliptic, glabrous, entire; leaf axils glabrous or tufted with hairs. Inflorescences terminal or axillary, paniculately or umbellately disposed cymes, often fascicled in leaf axils or defoliated nodes; flowers 4-5- (6-) merous; hypanthium campanulate or ovoid, glabrous or pubescent, rim very shortly dentate or truncate; petals thin, white or pink; stamens twice as many as petals, equal or unequal in size; filaments glabrous, flattened; anthers linear-lanceolate or linear-oblong, connective hardly produced, dorsally short spurred, ventrally with a pair of short appendages; ovary 4-5- (6-) celled; extraovarian chambers generally extending to the middle of the ovary; style terete, glabrous; stigma minute, punctiform or minutely capitate. Fruit a berry, globose to subglobose; pericarp thick or thin; seeds few to many, minute, ovoid, testa smooth or finely reticulate. Species Alliancea Neither Blume's nor Bakhuizen van den Brink's infrageneric classification was found suitable for characterizing the diverse species alliances in Borneo. I have therefore outlined the following 12 informally designated species groups which offer a convenient reference for further study and comparison (Table 1). While most species alliances are defined by comparatively trivial characters, it is hoped that such cases are here so grouped that the ultimate solution of their relationship will be facilitated. l8 ucoomonsm \maouneaw uCoomonsm ucoomonsm msouneaw mdounmaw msounmam msounwam msounwaw msounmaw msounmam msounmaw unopnmaw mMDHHHmm> measuoaoaw no moHoHommm hueaaaxe moaoacmn Hocaauou amaze HmcHEuou no hueaaaxe mosho Heaaeuou awake mo moaowommm humaafixe amaze mo moaowommm kumaawxe panama no mumuaaom moaho EuomHHHones Ho moaowcem hueaawxm mosho Buomwaaona: mumaafixo mosho Euomwaaones humaawxm mmamo Euowwaaones humHHme moHoHcma Hungaumu mozmommmodmzH ouecuouae ouwmommo euwmoaao oufimomao ouamoamo ouamommo ouamomao enamoaao ouwmoamo enamoamo ooauons voHHOLB umaoa ankx .ououmu QUQHQU GUGHGU Gumhmu unaswcmucmsv no unaswcwucmSUASm OUOHOU OUOHGU GUMkQU GUQHQD QUGHQU QUGHGU unaswceuvedv Vx¢HOAA>mm ZOHHomm mmOMU swam mnaunm mewmoouo no wGHnEHHo manuSm waHnEHHo madman oaumzawmo manunm Hmfiuumouuou HO mnsufim madHSm unsucm mflduSm owumnmwmo owuxsawam oauhnmwmo ouuhnafiao ofiumamwmo moon» Hanan uo mnsufim owuhsawmo mnsufim wanEHHo manage oauhcmwmo manunm Hmfiuumeuuou HO owuhnaaao HHm<= aaaaamamauan .aomm .NH mflmmmmmmmflmmmw .8 .HH egg 2': .oH mammwaawqammm . :I ox' NUCGHSUUSW . 2| «5 mwaMdMflMMNa .fl .5 mwloflflnilmflfid mflflmwflmmde . 2| .5 HHCMEWoA z] a' wumqwmm . 2| «i mHHHcHnoz .uoom .N manuaqmaa .m .H moz