A revision of the Ochnaceae of the Indo-Pacific area A. Kanis Contents Summary 1 Introduction 2 Subdivision and interrelations 3 Historical survey 3 Proposed subdivision 5 Morphological remarks 8 The vegetative parts 8 The inflorescence 8 The flower 10 Fruit and seed 12 Anatomical characters 12 The pollen 13 Geographical remarks 15 Summary of data 15 Discussion 17 18 Remarks on the presentation of data Family Ochnaceae 20 Subfamily 1. Ochnoideae 21 Tribe 1. Ochneae 22 61 Subfamily 2. Sauvagesoideae Tribe 2. Euthemideae 62 Tribe 66 3. Sauvagesieae 80 Excluded taxa Index 80 Summary This taxonomic revision includes all Ochnaceae from South and Southeast Asia, Malesia, Australia, and Pacific Islands. much wider is than in most works: in the A species concept applied previous 20 species listed. A few have 10 genera are accepted for the area treated, whereas 199 specific synonyms are taxa level which Kanis is been accepted at infra specific among Brackenridgea palustris ssp. kjellbergii new. overlooked in the and that the traditional It is shown that some old specific names have been past concepts have been in accordance with the All used for of some species not original concepts. names, currently L. accepted here, L. and G. serrata Asiatic species of Ochna and Gomphia Schreb. are no longer O. jabotapita for (Gaertn.) Kanis being the correct names the respective type species. O. fascicularis Blanco is made the of distinct section Notochnella Kanis in the Brackenridgea A. Gray. type a (v. Tiegh.) genus of the is and a subdivision of the Ochnaceae is A short history taxonomy given partly new suprageneric Two the Ochnoideae the tribes Ochneae and subsequently proposed. subfamilies are recognised: comprising Elvasieae Rchb., and the Sauvagesoideae Lindl. including the tribes Sauvagesieae, Euthemideae Planch., and in the subtribes Ochninae and Ouratinae (v. Tiegh.) Lophireae Rchb. The Ochneae are newly subdivided * Kanis, and the Sauvagesieae in the subtribes Sauvagesinae and Luxemburginae (Planch.) Kanis. in about characters the family, including some new Some general remarks are made morphological the is made towards better ofthe inflorescence characters of pollen. An attempt a understanding types. from Africa into Asia. Other in South- It is assumed that the genera Ochna and Gomphia migrated genera of east Asia, Malesia, Australia, and the Pacific Islands are regarded as long established, because the more less relict-like oftheir or nature areas. 1 2 BLUMEA VOL. XVI, No. i, 1968 Introduction The revision presented here was started as a preliminary study for a contribution to the FloraMalesiana.After completion of a first manuscript on the subfamilySauvagesoideae, extend it soon proved necessary to the studies in the Ochnoideae over the boundaries of Malesian of latter the area. Certain genera the subfamily are only poorly represented in Malesia, which made comparitive studies in related species from adjacent areas highly desirable. that A second, but not less important motive for this revision was the situation no the Ochnaceae critical monographic work including of the Indo-Pacific area was written after those by De Candolle (1811) andPlanchon (1846—47). The work of Van Tieghem been (1902—07) has merit in certain aspects, but his taxonomic treatment has not because of his Floras and accepted unusually narrow species concept. in Asia, Australia, the Pacific area could only give names according to local traditions. treated the and of South and Southeast The area covers tropical subtropical parts continental Asia, Malesia, Australia, and the Pacific Islands. This means that the natural and boundaries of the Ochnaceae are reached in the North (Himalayan Mts., Kwangsi, marks the the Hainan) and the South (NE. Queensland). Fiji eastern boundary, as it is remote island where Old World whereas most group an genus is found, neotropical of Pacific Islands. genera this family do not occur in the limits this chosen wide The western of revision are for practical reasons. There is a gap and of between the species in Indiaand Ceylon on the one hand those the SW. Arabian the other. the Peninsula, Africa, Madagascar, and the Mascarenes on However, two of treated here African and there is a rather close genera are mainly apparently relationship between certain species in continental Asia and others in Africa and Madagascar. I have refrained from analysing African material, since thorough comparitive studies could of this the easily have doubled the time necessary for the preparation paper. Besides, African Ochnaceae have been revised more often than their eastern relatives and presently the of less studies other botanists. they are subject more or parallel by and have been Some brief, general chapters on taxonomy, morphology, geography added of the as an attempt towards a better understanding family as a whole, though had almost with special reference to the taxa treated. For data on other taxa I to rely this Ochnaceae completely on literature. I hope that will facilitate comparisons with from the African region and the Neotropics. The of this made a from the Netherlands completion paper was possible by grant Foundation for the Advancement ofTropical Research (WOTRO) and by an allowance herbaria in London and Paris. This substantial is for a study tour to support gratefully acknowledged here. I H. director of the am very grateful to Professor Dr. J. Lam, formerly Rijksherbarium, who first the of and advised me to extend encouraged my steps on path plant taxonomy revision. his Professor my studies to this regional I am very much indebted to successor, first Dr. C. G. G. J. van Steenis, who made it possible for me to realize this plan, as his later as a worker. with all facilities from assistant, guest He kindly provided me possible much of his the Rijksherbarium and the Flora Malesiana Foundation and he spent valuablt time in reading and discussing the final draft of the manuscript. I should like thank to both scientists also for their major contributions to my general education as a botanist. den Dr. R. C. Bakhuizen van Brink Jr., Dr. W. A. van Heel, Dr. C. Kalkman, Dr. P. W. Leenhouts, and Dr. H. Sleumer, staff members of the Rijksherbarium, gave A. Kanis: A revision of the Ochnaceae of the Indo-Pacific Area 3 miscellaneous advices during my work, for which I want to thank them very much. indebted at the 1 am greatly to Mr. J. Muller, presently palynologist Rijksherbarium, made of in the of the who a preliminary inventory pollen types Ochnaceae, especially his taxa treated here, kindly putting results at my disposal. Gratefully acknowledged assistance was received from Miss E. van Nieuwkoop, who the from typed most of the final manuscript and compiled index, my wife, who also of different and from C. Marks and Mr. E. who typed part it in stages, Mr. L. Vijsma, prepared the drawings for this paper. The directors of the institutes mentioned belowkindly permitted examination of their collections. Their cooperation is gratefully acknowledged here. I have personally visited the institutes marked with an asterisk*. A Arnold Arboretum, Cambridge, Mass., U.S.A. BKF* The Forest Herbarium, Bangkok, Thailand. BM* British Museum (Natural History), London, England, U.K. BO Herbarium Bogoriense, Bogor, Indonesia. BRI Botanic Museum and Herbarium, Brisbane, Qld., Australia. C Botanical Museum and Herbarium, Copenhagen, Denmark. CAL Central National Herbarium, Howrah, Calcutta, India. DD Forest Research Institute, Dehra Dun, India. E Herbarium of the Royal Botanic Garden, Edinburgh, Scotland, U.K. FI Herbarium Universitatis Florentinae, Florence, Italy. G Conservatoire et Jardin Botaniques, Geneva, Switzerland. GH Gray Herbarium of Harvard University, Cambridge, Mass., U.S.A. GL Department of Botany, University ofGlasgow, Scotland, U.K. K* Herbarium of the Royal Botanic Gardens, Kew, England,U.K. KEP* Forest Research Institute,Kepong, Malaya, Malaysia. KYO Departmentof Botany, Kyoto University, Japan. L* Rijksherbarium, Leiden, Netherlands. P* Museum National d'Histoire Naturelle, Paris, France. S Naturhistoriska Riksmuseum, Stockholm, Sweden. SAN* Forest Department, Sandakan, Sabah, Malaysia. SAR* Sarawak Museum and Forest Department,Kuching, Sarawak, Malaysia. SING* Herbarium of the Botanic Gardens, Singapore. U* Botanical Museum and Herbarium, Utrecht, Netherlands. SUBDIVISION AND INTERRELATIONS Historical survey 1811 The Ochnaceae were first described as a family by A.P. de Candolle in \ but it until lasted 1895 that a more unanimously accepted circumscription was given by Gilg. such natural It is not surprising, however, that this family was not recognized as in early systems. The androecium and especially the gynoecium in the respective tribes show at resemblance. could first sight remarkable differences rather than a general Nonetheless, it eventually be demonstrated that the flowers of the Ochnaceae have much in common fundamentally, whereas anatomical and palynological evidence also indicates true inter- relationships. In De Candolle's concept, the family Ochnaceae comprised about what is treated here as the subfamily Ochnoideae. Initially (1811) he added a tribe Simaroubeae which he Ochnaceae distinguished later on as a separate family (1824). A close relationship between several authors. Reichenbach andSimaroubaceae was accepted long after by (1841) even accepted the Ochnoideae as a subtribe of a tribe Simaroubeae in the family Rutaceae. l taxonomic ) References to publicationsin this chapter are given more extensively in the enumeration, mostly under the family, sometimes under relevant lower taxa. BLUMEA VOL. No. 4 XVI, i, 1968 According to later opinions, this was based on analogy rather than on homology of certain floral characters. The genusSauvagesia L. was regarded by De Candolle (1824) as belonging to a distinct tribe of the of Violaceae, because its 3-carpelled, 1-celled ovary, developing into a 3- valved with seeds. the capsule many However, inSauvagesia capsules are opening along of the whereas Violaceae the fruits the margins carpels, in the are dehiscing along the authors medians. His view was supported by several like Meisner (1836), Bentham & and Hooker (1862), Le Maout&Decaisne (1868), Baillon (1873).