Myristicaceae

BLUMEA 25 (1979) 321 -478 New account of the genus Knema (Myristicaceae) W.J.J.O. de Wilde Rijksherbarium. Leiden, The Netherlands Contents Summary 321 Introduction 322 Acknowledgements 323 323 The characters in Knema 324 (1—3) Vegetative characters (4-9) Generative characters 326 330 The series in Knema: considerations and comparison with Sinclair's subdivision Keys to the series and species 338 339 (1) General key, mainly based on male flowering specimens 352 (2 — 6) Regional keys to female flowering or fruiting specimens Enumeration and descriptions of the species 365 Index 475 Summary 12 A tentative subdivision ofKnema into series, containing 83 species. The series and species aremainly defined by the shape of the mature male flower bud (perianth) and the androecium, with in addition various vegetative characters primarilyconcerning the tomentum of the apical part of the twigs, of the lower leaf surface, and of the flowers. This subdivision into series is partly fairly differing from the division into Sinclair in Gard. Bull. 16 and 18 which groups poposed by Sing. (1958) (1961), was mainly of and these latter characters based on characters the female flowers, mainly style stigma; are presently regarded as of less taxonomic significance. A in Knema is The in mainland SE. survey ofthe characters regarded asimportant given. genus occurs Asia and Malesia, from India to western New Guinea; not in Ceylon. Of the 83 species, 63 occur in Malesia, and 30 in mainland SE. Asia excluding Malaya and Singapore; 20 species are restricted to that 10 of into the moist latter area, and most of the remaining species are Malesian origin, just penetrating forest in Peninsular Thailand. Most in of which 22 endemic. evergreen species occur Borneo, 39, are A generalkey to all species of Knema, inside and outside Malesia, based mainly on male flower- and on vegetative characters, is presented. For female flowering and fruiting material five separate regional and four the Malesian keys, onefor mainland SE. Asia (excl. Malaya and Singapore), keys covering area, also using in addition vegetative characters, is presented. As compared to Sinclair’s monograph(1961), in which 37 species, 14 varieties, and 1 form were accepted, several species have presently been taken out of and the of of the synonymy re-installed, necessitating redescription many species. Many newly proposed species appeared to be not yet or insufficiently collected at the time of Sinclair's treatment. Description and discussion of the 12newly proposed series. Enumeration and treatment ofall species accepted. Several species have been subdivided into subspecies, varieties, or forms. Besides the 12 new 8 in addition series, 25 new species, new subspecies, 3 new varieties, and 3 new forms are described, and there are several proposals ofa new status of already existing taxa. For all new names and combinations, see the index. All extra-Malesian taxa are fully treated, providing for synonymy, typification, description, notes, and enumeration of examined specimens; ofthe Malesian species usually merely the full ofthese synonymy, types, some notes, and examined specimensare presented,as descriptions species will in Flora Malesiana. An index of all scientific Knema is the end. appear names pertainingto given at 322 BLUMEA - VOL. 25, No. 2, 1979 Introduction is Besides Myristica, Horsfieldia, and Gymnacranthera, the genus Knema one of the four Asian genera of Myristicaceae. It is distributed from S. and NE„ India through SE. Asia to S. China and Indo-China, and throughout Malesia, east to W. New Guinea (Vogelkop Peninsula). Its greatest species density is in Borneo, and there are smaller centres of species development in the Malay Peninsula and Indo- China. While the revisions of the the late Sinclair for working up genus Knema by J. the treatment ofthe family Myristicaceae in the Flora Malesiana, it soon became clear that reconsideration of again a personal the whole genus was necessary. Sinclair's work on Knema was published in two papers, one dealing with the family Myristi- caceae for Malaya, published in Gard. Bull. Sing. 16 (1958) 254 — 332, and an additional with the whole in and outside in publication dealing genus, Malaysia, Gard. Bull. Sing. 18 (1961) 102 - 327; this latter also being a precursor for his then anticipated treatment in the Flora Malesiana. Both works contain very much information the offer taxonomic of the on genus; they a subdivision, a survey criteria for the delimitation of the species, several keys, and descriptions and of drawings species, including field characters. Useful general chapters on mor- phology can be found especially in the publication of 1958. These works greatly enriched our knowledge of the genus since Warburg's monograph of the Myristi- This caceae (1897). latterwas, at least for Knema unsatisfactory, as it obviously was , based on too few specimens. King's account for former British India, in Ann. Roy. Bot. Gard. Calcutta 3 (1891) 284 — 331, where the genusstill is treated as sections of be useful for the of from Myristica, appeared to very interpretation several species that area. Since 1961 the amount of unidentified material of Knema from Malesia in the Leiden herbarium had considerably augmented. One of the difficultiesencountered with the determination of this material with Sinclair's keys, as well as with the of incorporation the rather numerous newly discovered facts into Sinclair's scientific that Sinclair's founded the framework, was system was primarily on structure of the style and stigma, present only in the pistillate flowering specimens of the dioecious it that the criterion for genus. Moreover, appeared main his classification, namely whether the stigma is few- or many-lobed, proved to be in many cases variable and much in too overlapping, also specimens studied or identified by Sinclair himself. This led me to investigate the flowers of both sexes still more intensively than Sinclair did. It was found that in the staminate flowers several fairly distinct types could be distinguished, based on the general shape and architectureof the perianth and the androecium, characters that were underestimated by Sinclair. In addition, several other new findings proved to be of taxonomic importance, such as the absence ofsmall non-traumatic warts presence or cork on the lower leaf surface, and a more accurate ofthe and ofthe tomentum of description composition appearance the flowers, the twig apices, and the lower leaf surface, as well as of characters introduced the size of the in already by Warburg, e.g. perianth expressed the diameterof the mature male flower buds. With the introductionofthese characters it appeared that several of the difficult, i.e. rather obscure variable taxa defined or very as by Sinclair, could be better understood by accepting many more species. These hadpartly to be newly described W. J. J. O. de Wilde: Knema 323 whereas in several I to the conclusion that or taken out of synonymy, cases came rank could better be treated distinct taxa accepted by Sinclair in an infraspecific as material at for the of species. In some cases, the my disposal recognition a new in Sinclair's time collected. species was not yet, or merely very incompletely, Sinclair in the All species names accepted by are retained present study, though redefined. the these had, as a matter ofcourse, sometimesto be fundamentally Thus, number of species in Knema has increased from 37 (with 14 varietiesand 1 form) as in accepted by Sinclair (1961), to as much as 83 in the present survey. Warburg, varieties in 1897, recognized some 39 species and 18 Knema. short account I have, for most a commentary as In my present species, presented to the differences with Sinclair's opinion, and for the new- and re-installed species explained why these were not recognized by him. the Only the extra-malesian taxa are fully described in the present paper, as Malesian species will be completely dealt with in the Flora Malesiana treatment. ACKNOWLEDGEMENTS of British My gratitudeis herewith expressed to the directors and keepers of the herbaria the Museum ofNatural History (London),the Royal Botanic Gardens, Kew, and ofthe Natural History Museums at and sending me selected specimens on loan. Paris and Utrecht for enablingme to work in their collections for The herbaria at Singapore,Bogor, Aarhus, and Copenhagenare acknowledged sending me on loan much of the unnamed specimens there. The herbarium of the Arnold Arboretum, Harvard University, generously provided the loan of a type specimen. The visit to the Paris herbarium was made possible by a grant from the Netherlands Organization for Pure Scientific Research (Z.W.O.), the Netherlands. Dr. R. C. Bakhuizen van den Brink and Dr. H. O. Sleumer generously provided for most of the translation of the diagnoses of the new taxa into Latin. The drawings of 18 species were prepared by Miss Ruth van Crevel and Mr. J. van Os of the Rijksherbarium. THE CHARACTERS IN KNEMA delimitationand classifi- Much information on the characters to be used in the in cationin Myristicaceae in general, inch Knema, is presented by Sinclair, especially his publication of 1958 dealing with the representatives of the family in Malaya. characters have been and it However, as some valuable additional found, as appeared that certain features should be estimated somewhat differently as compared to Sinclair's practice, I have enumerated below those characters which are useful and which have served in the delimitation of the considered by me as species accepted in the present account. Someofthese characters should be judged as being important enough to warrant subdivision of the into these coincide with the a genus series; as a whole, larger divisions made in the general key to the male specimens. Also, I feel that these have and that most species in a characters may some evolutionary significance, With the series generally will have mutually a closer alliance.

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