Notes on Malesian Author(s): B. Verdcourt Source: Kew Bulletin, Vol. 40, No. 1 (1985), pp. 213-224+ix Published by: Springer on behalf of Royal Botanic Gardens, Kew Stable URL: http://www.jstor.org/stable/4108497 Accessed: 03/12/2010 05:09

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http://www.jstor.org Notes on Malesian Chloranthaceae

B. VERDCOURT

Summary. subsessilis is described as new and three new combinations are made in and Chloranthus. A summary of the literature dealing with the systematic position of the family is given. Whilst preparing an account of the Chloranthaceaefor Flora Malesiana some difficulty was experienced in delimiting the taxa of Ascarina. Examination of the extensive material at Leiden showed that some specimens from the high- land area of Papua New Guinea were distinctly different in foliage from A. philippinensis C. B. Rob. with which they had been confused; these have been ascribed to a distinct species which is described below.

Ascarina subsessilis Verdc. sp. nov. affinis A. philippinensis C. B. Rob. sed foliis subsessilibus carnosioribus satis distincta. Typus: Papua New Guinea, Western Highlands, Hoogland & Schodde 7331 (holotypus L; isotypi CANB, E, K). Arbor glabra, dioica, 15-24 m alta, cortice fissurato saturate brunneo vel pallide griseo-brunneo obtecta; ramuli longitudinaliter in siccitate irregu- lariter costati nodis aphyllis supra ipsos foliiferos praesentibus. Folia saturate viridia, elliptica, oblongo-elliptica, vel raro anguste obovata, 5-14 cm longa, 3-9 cm lata, apice rotundata, basi angustata vel rotundata, late cuneata usque subcordata, margine haud profunde crenata, subcarnosa, supra in siccitate minute rugulosa, subsessilia, petiolo crasso 2 mm longo; nervi laterales ? 30, subpatentes, utrinque in siccitate prominentes; costa laminam interdum inaequaliter dividens; stipulae verae minutae, + 1 mm longae; vestigiae foli- orum ad nodas aphyllas 0-8-1-5 mm longae, vagina 2-6 mm longa in siccitate rugulosa. Inflorescentiaeterminales, ramosae et interdum simplices in axillis paenultimis positae, 1-5-4 cm longae; rami primarii 3-5 numero, 5-7-divisi, ramulis crassiusculis 0-5-2-5(-3) cm longis in siccitate costatis, bracteis superi- oribus lanceolatis longitudinaliter plicatis 4-5 mm longis, bracteis inferioribus vestigias foliorum supra commemoratas simulantibus, bracteolis deltoideis vix 1 mm longis. Floresfeminei haud visi. Inflorescentiaemasculinae 2 cm longae, basi 3-ramosae; ramus primarius ramulosus, ramulis + 5 mm longis. Flores masculini 2-staminati, antheris 2-3 mm longis. Fructus viridi-albi, probabiliter in maturi- tate + nigri, 3 mm longi, 1-5 mm lati, congesti, exocarpio in siccitate forte rugoso; endocarpia immatura (?) 1 mm longa, laevia. (Fig. 1, Map 1). PAPUA NEW GUINEA. Morobe Distr.: Lae subdistr., Kasanombe, road to Momsalom village, mountain-ridge rain-forest, 1830 m, 29 Aug. 1973, Katik & Taho NGF 37914 (A, BRI, CANB, L, LAE):- tree 15 m tall, bole ? 8 m [sphalm 80 m], d.b.h. 24 cm, bark dark brown, under-bark green, inner bark cream, wood straw, leaves dull green, fruit green & Sattelberg, Ogeramnang Forest, 1740 m, 25 Jan. 1937, Clemens5123 (B, L):- illegible note. Southern Highlands

Acceptedfor publicationFebruary 1984 213 214 KEW BULLETIN VOL. 40(1)

FIG. 1. Ascarinasubsessilis. A male flowering branchlet x Z; B stipules x 4; C portion of male inflorescence x 8; D base of anthers showing bract x 16; E T.S. anther x 16; F female fruiting branchlet x 2; G portion of young female inflorescence x 8; H portion of older female inflorescence x 8; J L.S. female flower x 16; K fruits x 6; A-E from Saunders1034; F, G & K from Hoogland& Schodde7331; H & J from WomersleyNGF 14273. Drawn by Mrs M. E. Church. NOTES ON MALESIAN CHLORANTHACEAE 215

00

-0

I I I 1400 1450 1500 MAP 1. Distribution of Ascarina subsessilis.

Distr.: Mendi subdistr., 32 km from Mendi, Tambul road, Mimimbipl, 6' 00' S, 1430 40' E, montane mixed forest, 2300 m, 2 Sept. 1972, WomersleyLAE 55316 (BRI, CANB, L, LAE):- canopy tree to 20 m, bole 10 m, d.b.h. 20 cm, outer bark pale grey-brown, vertically fissured with long open pustular lines, underbark pale brown with darker fibres, wood pale brown, rays prominent. Western Highlands Distr.: Hagen subdistr., Tambil-Tomba road, Nebilyer divide, lower montane rain-forest, 2400 m, 17 July 1957, Robbins 469 (CANB, L):- tree mostly 15 m but mature specimens in forest up to 24 m, fairly com- mon, vern. name 'porn' (Hagen); Wabag subdistr., Sirunke, 2580 m, 19 July 1962, WomersleyNGF 14273 (L, LAE):- tree 15 m, leaves dark green, rather fleshy, flowers and fruits greenish white; same subdistrict, top of ridge N of Pombabus, lower montane forest, 42' slope, drainage fair, aspect SW, shade strong, brown forest soil, 267 m, 11 May 1965, Flenley ANU 2779 (A, BRI, CANB, K, L, LAE, US):- tree to 5 m, d.b.h. 8 cm, bark light brown, inner bark white, wood light brown, occasional, vern. name 'wabaliok' (Enga); Laigam subdistr., Sirunki-Laigam road, 6-4 km S of Lake Iviva, lower montane rain- forest, 2460 m, 23 Aug. 1960, Saunders 1034 (CANB, L):- tree 15 m tall, girth 75 cm, vern. name Wabilak (Enga) & Lagaip valley, near Kepilam Village, in tall forest at least partly secondary on limestone ridge, 2400 m, 5 Aug. 1960, Hoogland & Schodde 7331 (holotype L; isotypes CANB, E, K):- tree 16 m tall, 6 m bole, 35 cm diam., fruits green, vern. name 'wabaljok' (Enga, Kepilam). Eastern Highlands Distr.: Goroka subdistr., W of Fatima R., Marafunga Mill, 6? 05' S, 1450 15' E, primary montane rain-forest, 2600 m, 25 Nov. 1970, Grubb 216 KEW BULLETIN VOL. 40(1) & Edwards 59 (A, L, LAE):- tree to 23 m d.b.h. 195 cm (sic) & Lufa sub- province, W slopes of Mt Michael, 6' 27' S, 145' 19' E, mostly primary rain-forest but showing much evidence of disturbance by man, 3200 m, 13June 1979, Sohmeret al. LAE 75435 (A, BRI, CANB, E, K, L, LAE, UPNG):- tree 15-20 m tall, bole 10 m, d.b.h. 20 cm, immature fruit green.

Sarcandra glabra (Thunb.) Nakai, Fl. Sylv. Koreana 18: 17 (1930). Type: , Thunberg (holotype UPS). Bladhia glabra Thunb., Trans. Linn. Soc. Lond. 2: 321 (1794).

subsp. glabra Male structure with non-antheriferous basal part well developed. DISTRIBUTION. Japan, N & C , .

subsp. brachystachys (Blume) Verdc. stat. & comb. nov. Ascarina serrata Blume, Enum. P1. Jav. 1: 80 (1827). Type: Javae montibus altioribus, Blume, Reinwardt (? syntypes L). Chloranthusbrachystachys Blume, Fl. Jav., Chloranthaceae: 13, t.2 (1829). Type: based on Ascarina serrata Blume (non Chloranthusserratus (Thunb.) Roem. & Schultes). Chloranthushainanensis Pei in Sinensia 6: 674 (1935). China, Hainan, Lin Fa Shan, Tsang 317 (isolectotype K). Sarcandra hainanensis (Pei) Swamy & Bailey in J. Arn. Arb. 31: 128, figs. 4, 15-17 (1950). Male structure with non-antheriferous part very reduced or absent, i.e. the anther cell more or less as long as the structure. DISTRIBUTION.S China to NE and throughout Malesia from Malaya to Papua New Guinea. When Swamy & Bailey (1950) dealt exhaustively with the anatomy of Sarcandra and reinstated the they treated Chloranthusbrachystachys as a synonym of Sarcandraglabra but kept up S. hainanensis (Pei) Swamy & Bailey stating that it had discoid male structures with the anther cells as long as or almost as long as the whole structure and also gave a number of other charac- ters, including anatomical, of little actual taxonomic significance. Typical S. glabra from Japan and northern China has the non-antheriferous part of the male structure quite well-developed. Dr Ohashi has kindly informed me that an examination of Japanese material shows little variation in the relative lengths of the anther-cells and the non-antheriferous part although there is a gradation in size from north to south. Chinese material I have examined supports this. All tropical material of S. glabra has the structure very similar to that depicted for S. hainanensisand I consider subspecies should be recognized. The anomaly of having a species widespread in Malesia and Asia save in Hainan is therefore eliminated; rather there is a tropical subspecies extending to S China and a more northern subspecies. The occurrence of some slightly intermediate specimens in northwest India and probably in China suggests that specific rank is not correct; if it were so decided the epithet serratawould have to be employed. It is clear from the Flora Yunnanica (1: 21 (1977)) that NOTES ON MALESIAN CHLORANTHACEAE 217

both S. glabra and S. hainanensisare considered to occur there but C. brachystachys is still placed in the synonymy of the former.

var. brachystachys Fruits orange or red.

var. melanocarpa (Ridley) Verdc.comb. nov. Chloranthusbrachstachys Bl. var. melanocarpaRidley in J. Mal. Br. Roy. As. Soc. 87: 89 (1923). Type: Sumatra, Ridley (holotype SING). Fruits black.

SUMATRA. Berastagi, West Hill, Ridley (holotype SING; isotypes K). Gunung Batu Lopang, 10 km ESE of Propar (Lake Toba), 1400-1500 m, de Wilde & de Wilde-Duyfjes 13508 (L). Throughout its range Sarcandraglabra is described as having an essentially red fruit and I had looked on Ridley's specimen as a chance variation. A second specimen with black fruits from the same general area of Sumatra indicates that populations exist in restricted areas and I have decided to keep up Ridley's varieties. The additional leaf-characters he mentions do not hold.

The following new combination is also needed. Reading through my pre- liminary manuscript Dr C. G. G. J. van Steenis noted what had escaped my attention; that I had placed an earlier name in the synonymy of a later one.

Chloranthus erectus (Buch.-Ham.) Verdc. comb. nov. Cryphaeaerecta Buch.-Ham. in Edinb. J. Sci. 2: 11, t. 2 (1825). Type: India, Goyalpara, Buchanan-Hamilton98 (holotype E!; photo. K). Chloranthusofficinalis Blume, Enum. P1. Jav. 1: 79 (1827) & Fl. Jav., Chlo- ranthaceae: 10, t. 1 (1829). Type: Java, Blume (syntypes L!). ? C. elatior R.Br. ex Sims, Bot. Mag.: 48, sub t. 2190 (1820) nom. nud. Type: not found. ? C. elatior Link, Enum. P1. Berol. 1: 140 (1821). Type: specimen grown at Berlin which did not flower (holotype Bt). Swamy and Bailey synonymised Cryphaeaerecta with Sarcandraglabra being influenced by the poor figure and very misleading description. I have, how- ever, borrowed the type and there is no doubt it is identical with Chloranthus officinalis.Very unfortunately the earlier C. erectusof Sweet (Hort. Surburban. Lond.: 28 (1818)) is a nomen nudum-the single word upright is merely to supply an English name and in any case would scarcely qualify as a descrip- tion. The name C. elatior has come into widespread use in the past 20 years but Robert Brown's name elatior is not validly published since the character he mentions is used to distinguish elatior and another species from a third species. There is no description of elatior as such. Link describes a sterile specimen said to have been received from an English garden and the information given is not adequate to show that Link's specimen was conspecific with C. ofJicinalis;he does not mention Robert Brown so the citation C. elatior R.Br. ex Link fre- quently used is not correct. Link does suggest his C. elatior might be the same as C. erectusSweet. 218 KEW BULLETIN VOL. 40(1)

THE POSITION OF THE CHLORANTHACEAE: A SURVEY OF RECENT IDEAS

For a small family the Chloranthaceaewith only four extant genera Chloranthus Sw., Sarcandra Gardner, Ascarina J. R. & G. Forster and HedyosmumSw. has engendered an extraordinarily large literature. Most collectors and those engaged in routine naming might well agree with a remark I found on a specimen of Chloranthusfortunei(A. Gray) Solms collected by Reginald Farrer on 7 May 1914 who dismissed it in two words 'ugly thing'; maybe, but the lack of beauty in the family is certainly compensated by its intense interest from an evolutionary point of view. Although Swamy (1953) speaks of the establishment of the family Chlo- ranthaceaeby Bentham & Hooker f. (1880) it actually dates from R. Brown (1821) ex Lindley (1821). Early placings were wide of the mark. Jussieu (1808) placed Chloranthusin the Loranthaceae,Sprengel (1817) in the Caprifoliaceaeand Brongniart (1850) included the family in his Santalineae. Cordemoy considered the family allied to the Ceratophyllaceaeand Van Tieghem (1884) put them near the Casuarinaceae,Ceratophyllaceae and Platanaceae. Endlicher (1837) and Lindley (1846) placed them in the Piperales and his view has been supported by numerous authors to the present day and is, in my opinion, at least close to the correct affinity. Bentham & Hooker (1880, 1883) included the Chloranthaceae in the series Micrembryeae of the order Monochlamydeae where it comes next to the Piperaceae, Myristicaceaeand Monimiaceae. At that time Circaeasterwas still included in the family but this is now referred to a separate family Cir- caeasteraceae;it differs in many ways, particularly the absence of secretory cells. Engler (1894) agreed with the, by then, conventional view associating Chlo- ranthaceaewith Saururaceaeand Piperaceae within an order Chloranthales but Johnson (1905) did not think there was a close relationship with Piperales. Armour (1906) carried out detailed morphological studies of Chloranthuschinen- sis (untraced name), C. officinalis(= C. erectus)and C. brachystachys(= Sarcandra glabra) and found that the ovule of Saururus is intermediate between that of Chloranthusand the Piperaceae since in it the mother-cell of the embryo-sac undergoes division while the subsequent development of the seed resembles Piper. In Chloranthusthe mature embryo-sac and later stages of development present no peculiar features. The most probable conclusions from the floral structure appear to be that the Chloranthaceaeis a group of the Piperales presenting in some points and, especially in the structure of the ovule, primitive characters in common with the majority of the Archichlamydeae while in other respects special modifications of the flowers are shown. Hallier f. (1912) placed the Chloranthaceaewith Annonaceae,Magnoliaceae and Laurales in the Annonales under his broad grouping Annonophylae which says scarcely more than that it is a primitive family. Thierry (1912) concluded that Chloranthaceaeand Saururaceaehad more anatomical characters in common with each other than either had with the Piperaceae.Swamy (1953) discussed the affinity of the family in detail and gave reasons for not associating it closely with the Piperales. Apart from the vesselless xylem of Sarcandra the other genera have 'decidedly prim- itive cambium resulting in xylem of only slight specialisation whereas the xylem of Piperaceae,Saururaceae and Lactoridaceaeis more highly evolved.' Con- spicuously wide rays are not evidence of close affinity with Piperaceaesince they also occur in Lauraceae, Monimiaceae etc. His main objection to the usually proposed affinity with the Piperaceae is, however, based on nodal anatomy which is of unilacunar type in the Chloranthaceaein sharp contrast to the NOTES ON MALESIAN CHLORANTHACEAE 219 situation in the Piperaceaeand Saururaceaewhere it is typically multilacunar. He also claimed support from pollen stating 'the pollen of the Piperaceae and Saururaceaeappears to have undergone a particular trend of specialisation that has resulted in the formation of uniformly smaller sized grains with very faint or no sculpturing on the exine, and with a single germinal furrow. Although the pollen grains of Ascarina are monocolpate those of other genera show a polycolpate condition with a strong tendency towards the elimination of ger- minal furrows. This trend has culminated in the acolpate grains of Sarcandra and of some species of Hedyosmum;the exine in all genera exhibits a reticulate- pitted sculpturing of varying grades; the grains in general are relatively large'. He considered that the floral similarities between Chloranthaceaeand Piperaceae had been overemphasized. What embryological information there is indicates at least some differences between the families. 'The nearest relatives of the Chloranthaceae are those families of the ranalian affinities that possess mono- colpate pollen, ethereal oil cells and unilacunar nodes' which comprise Austro- baileyaceae,Trimeniaceae, Amborellaceae, Monimiaceae, Gomortegaceae,Hernandiaceae, Calycanthaceaeand Lactoridaceae.Endress (1971) also stated that the gynoecium structure lends support to the recent view of a close patristic relationship of the family with the Laurales-Monimiales. What may be termed the new mor- phologists have naturally paid some attention to such a primitive family. On Melville's gonophyll theory the male flowers of Chloranthusand Sarcandra are easily derived from androphylls by reduction; Sarcandrabeing similar to items 3-5 in the sequence showing primary evolution of a terminal androphyll (Melville 1963, fig. 65) save that Sarcandra has 2 vascular strands in what is usually termed the filament. In the same figure items 10, 18-27 showing the reduction of a complex androphyll to foliar stamens of classical theory include Chloranthus(24). Evolution of this laminar type is traced here from Hedycarya (Monimiaceae) through a sequence of intermediates formed by reduction ending with Doryphora and Nymphaea; from the intermediates, Chloranthusand Austro- baileya are easily derived. Meeuse, who has written extensively on the probably multiple rather than monophyletic origin of the Angiosperms, believed (1963) that there is a direct homology between the chlamydote ovules of the Chlamy- dospermae,Pentoxylales and other bennettitalean groups and the 1-ovuled gynoe- cia of many Piperales, Juglandales, Pandanales etc. He (1963, fig. 1) derived the Piperales in which he included the Chloranthaceaefrom part of the Ben- nettitales and possibly sharing common ancestry with the Gnetales, stating that the Chloranthaceaein particular exhibit certain primitive (gymnospermous or bennettitalean) characters namely the lack of vessels in the secondary xylem of Sarcandra,the presence of a narrow open channel leading from the stigmatic part to the ovary chamber in Chloranthus and Sarcandra, the orthotropous ovules, the reported occasional occurrence of an integument protruding from the ovary (Yoshida 1959; Meeuse 1963, fig. 3), embryosacs sometimes with 20-40 antipodal nuclei of Peperomia type reminiscent of the structure of the embryo sac in the Chlamydospermae, minute incompletely developed embryo at time of detachment of fruit, developing in the fallen seed before it germinates as in some Cycadales and Gnetum (Yoshida 1957, 1959a, b) and pollen grains similar to those of the Gnetales. In fact the lack of vessels is not shown by the Gnetales and in this respect Sarcandrais supposedly more primitive but I think the attribution of polarity to character-states is very often quite naive. The genetic bases are usually unknown and reversals might be much easier than seems plausible. Certainly the lack of vessels in Sarcandraand their presence in 220 KEW BULLETIN VOL. 40(1) Chloranthusis such a case. Sarcandra flowers could be derived from those of Chloranthusand the lack of vessels a reversion not a primitive state. Young (1981) has recently conducted a cladistic analysis of selected families of the Magnoliidae, Ranunculidae and Hamamelidae and shown that the most rea- sonable suggestion is to consider the absence of vessels in dicotyledons as a derived feature thus refuting the very widely accepted hypothesis that angio- sperms are primitively vesselless although Jeffrey & Cole already suggested otherwise in 1916. Meeuse supposed that if a Chloranthaceaturned up with consistently exposed integuments it would have to be referred to the Chlamydo- spermae but there are a multitude of other characters which are not shared. Elsewhere Meeuse has stated (1962) 'I regard the Piperales as practically inseparable from the Chlamydospermaebut do not think they are of high antiq- uity chiefly due to their exceptional eurypalyny'. Melville (personal commu- nication) considers it impossible that any of these groups had their origin in the Bennettitales but rather the Glossopteridae.Vijayaraghavan (1964) has made extensive studies of the embryology of Sarcandrachloranthoides Gardner (= S. irvingbaileyi Swamy) and decided that the sum total of the evidence from unilacunar nodal anatomy, reticulate exine sculpture, megasporogenesis (tet- rads), Polygonatum type embryo-sac, cellular endosperm, transverse first division of the zygote, absence of perisperm and presence of endosperm fails to support any relationship with Saururaceaeand Piperaceaeand would place it in the Ranales. Walker (in Beck 1976) in his classification of the Ranalean Complex used for expressing his palynological results, divides the Piperales into Chloranthineae,Lactoridineae and Piperinae the latter containing Saururaceaeand Piperaceae.Xi (1980) in another study of the pollen morphology of the Piperales confirmed the correctness of the retention of Chloranthaceaewithin that order but suggested they were sufficiently distinct to form a suborder considered the more primitive. He made a number of statements in support which seem over-simplified e.g. the suggestion that the exine is adopted to entomophily which is a condition more primitive than anemophily but considering the known fossil record of the insects this is doubtful if not quite untrue. Other supporting evidence claimed is the large size of the grains, the existence of Clavatipollenites(apparently indistinguishable from Ascarina) in the Early Cre- taceous and the vesselless xylem of Sarcandra and delayed development of its embryos. A. C. Smith (1973) in a roneoed sheet attached to reprints of his paper (?1972) stated 'the family Chloranthaceaeappears extremely distinct and in some respects more primitive than other elements of the Laurales. I would now suggest for it a separate order Chloranthales' but this cannot I think be held to constitute valid publication of the order. In the actual paper he included the family in the Laurales-Monimiinae between Calycanthaceaeand Lactoridaceaethus essentially following Swamy but the latter family he also placed in a separate order in the corrigenda sheet just mentioned. Smith (1976, 1981) repeated that the family 'is best assigned to its own order' and Leroy (1983) has validated it. Hutchinson (1973) and Cronquist (1968, 1981) retain Chloranthaceae in the Piperales but Takhtajan (1969) referred it to the Lau- rales. Patel's (1975) study of the wood anatomy of Ascarina lucida shows that there is little to support the views of Swamy and Takhtajan. Burger (1977) has also reviewed the evidence for the position of the family and pointed out that it shares with the Piperales the tendency for the stamens to become adnate to the back of the ovary, orthotropous ovules, sheathing leaf-bases, thickened nodes and certain features of stem anatomy. Moreover, he pointed out that NOTES ON MALESIAN CHLORANTHACEAE 221 Howard (1970) has demonstrated that the nodes are not modified unilacunar as indicated by Swamy but rather a modified trilacunar type, that the Chlor- anthaceaelack the flavonoid isorhamnetin so characteristic of the Laurales and chemically resemble the Piperales in many ways, that chromosome numbers suggest relationship rather with the Piperales and that Behnke (1975) has shown that S-type sieve-tube plastids occur in Chloranthaceae as in all investigated Piperales. Burger concluded that separate ordinal rank or prefer- ably subordinal rank within the Piperales are the best solutions. My own views based on mere 'gestalt' support the classical view of the Piperales but the rather extensive evidence against this view adduced by Swamy (1955) and Vijayaraghavan (1964), mostly ignored by others, suggests that Smith's erec- tion of a separate order is advisable but not very helpful in determining the true affinities. There does seem to be a more direct link with the Gymnosperms than is shown by other Angiosperms, relationships cropping up again and again in quite different types of characters. Burger (1977) has pointed out many often striking similarities between the Piperales and some Monocotyledons such as Aponogetonaceae,Potamogetonaceae and Araceaeranging over the wide spectrum of ecology, form, leaf-form, anatomy, inflorescences, flowers, pollen and chemistry and believes this cannot result from convergence from very different ancestors. He accepts the small simple flowers of the Chloranthaceaeas primitive rather than reduced and that the flowers ofLilaea (Juncaginaceae) may be primitively simple. Although many of the pistils of Lilaea have lost their subtending bracts and few bisexual flowers occur, the individual bisexual flower consisting of bract, stamen and simple pistil appear to be directly comparable and perhaps homologous with the flowers of some Chloranthaceae.There is no direct palaeontological evidence as yet but the similarity of Chloranthaceaepollen to Clavatipollenitesand the partic- ular similarity of another early Cretaceous pollen Asteropollis to that of some species of Hedyosmumsuggest the family has an ancient lineage explaining the persistence of some very primitive characters. The flowers can all be derived from an original plan of a single bract, 2 or 3 stamens and solitary pistil rather than by reduction from more complex spiral or whorled precursors-a view diametrically opposed to Leroy's theory of the male flower of Hedyosmum. Adaptation to wind-pollination has been given as evidence of the extreme reduction of the flowers and an objection to considering them primitive. Some Hedyosmumspecies are undoubtedly wind-pollinated but the forest-floor dwell- ing Sarcandra and Chloranthus cannot be wind-pollinated and although no nectar is produced it has been widely argued that insect-pollination began by pollen-collection; the aromatic smell so characteristic of e.g. C. spicatus is suggestive. Burger hypothesized that the flowers of the Saururaceae,Piperaceae, Lactoridaceae,Potamogetonaceae, Aponogetonaceae, Alismatales and Nymphaeales originated by condensations of simpler bisexual flowers similar to those of the Chloranthaceaeand Lilaeaceaeand that these families belong to a common almost prefloral stage before the Monocotyledons and Dicotyledons diverged. The floral vasculature of Lactorisstrongly supports Burger's theory. Melville's inter- pretation of three fused androgynophylls is similar. Burger explains the very prevalent trimery of monocotyledonous flowers by the fact that the simple flowers they derived from had an arrangement such as is prevalent in Chlor- anthus where the flowers are in opposite pairs or occasionally show a I 'phyllo' taxy. Condensation of more than one full cycle probably presented difficulties and accounts for the restriction. He makes the very plausible remark that it is 222 KEW BULLETIN VOL. 40(1) more reasonable for an additive process to result in almost constant trimery than that a reduction process should do so. Such fusion of female parts does actually occur in two Hedyosmumspecies where a usually solitary pistil is partly enclosed within a cup-like or tubular structure; but occasionally the tube will contain 2 or even 3 pistils easily interpreted as the fusion of 3 bracteate flowers with the usual loss of 2 pistils. Even if this is probably a recent happening it indicates a possibility of what may have occurred in the past in other genera. The problem of the origin of the perianth is accounted for by the derivation of one whorl from the bracts which protect the primitive flowers of Chloranthus and a subsequent doubling to improve protection and thence by modification to provide a whorl adapted to insect attraction. Evidence for this is given by the ontogeny of flowers in various Alismatales. Leroy (1981, 1983a, 1983b) has newly interpreted the male flowers of Hedyosmum.Formerly considered to be tightly packed bractless flowers reduced to a single anther he interprets the strobiloid arrangement of anthers as a single flower rather than an inflorescence. This cone-like arrangement coupled with monosulcate pollen of a type known from the Lower Cretaceous and anemophily suggests it could be close to the ancestral Angiosperm flower. This polystaminate Hedyosmumflower is perfectly homologous with the male gymnosperm cone. Ascarina which is insect-pollinated could have been evolved from a common ancestor by reduc- tion in the number of stamens. Chloranthus and Sarcandra are clearly of a different line but one can but guess at the diverse forms which have become extinct since the Cretaceous. This elegant and very plausible interpretation of the male flower of Hedyosmumhas been accepted. Dr P. Rudall of the Jodrell Laboratory, Kew kindly made sections through the floral axis of spirit material of Hedyosmumorientale collected by Mr B. L. Burtt in Borneo and kindly loaned by him. There are solitary traces to the anthers with no remnant of any others which might have been expected if they were individual reduced flowers although of course other traces could have completely disappeared. It is certainly clear that the now insignificant but still widespread family Chloranthaceaeis one of great importance in the study of primitive flowering .

REFERENCES

Armour, H. M. (1906). On the morphology of Chloranthus.New Phyt. 5: 49-55, pl. 3 & 4. Beck, C. B. (1976). Origin and early evolution of angiosperms. New York. Behnke, H.-D. (1975). The bases of Angiosperm phylogeny: ultra structure. Ann. Missouri Bot. Gard. 62: 647-663. Bentham, G. & Hooker, J. D. (1880 & 1883). Genera Plantarum 3 (133-135 & 1220-1221). London. Brongniart, A. (1850). Enumeration des genres de plantes Cultives ... Paris. Burger, W. C. (1977). The Piperales and the Monocots-alternate hypotheses for the origin of Monocotyledonous flowers. Bot. Review 43: 345-393 (very extensive bibliography). Cordemoy, C. J. de (1863). Monographe du groupe de Chloranthac'es. Adansonia 3: 280-310. Cronquist, A. (1968). The evolution and classification of flowering plants. New York. - (1981). An integrated system of classification of flowering plants. New York. NOTES ON MALESIAN CHLORANTHACEAE 223

Endlicher, S. (1836-40). Genera Plantarum secundum ordines naturales dis- posita. Vol. 1 (264 (1837)). Vienna. Endress, P. K. (1971). Bau der weiblichen Bluiten von Hedyosmummexicanum Cordemoy (Chloranthaceae). Engl. Bot. Jahrb. 91: 39-60. Engler, A. (1887). Natiurlichen Pflanzenfamilien 3(1): 12-14. Leipzig. Hallier, H. (1912). L'origine et le systeme phylktique des Angiospermes exposes A l'aide de leur arbre gne'alogique. Arch. Neerl. Sci. Exact. Nat. 11IB, 1: 146-241 (207-208). Howard, R. A. (1970). Some observations on the nodes of woody plants with special reference to the problem of the 'split-lateral' versus the 'common gap'. Bot. J. Linn. Soc. 63 Suppl. 1: 195-214. Hutchinson, J. (1973). Families of flowering plants. ed. 3: (513-519). Jeffrey, E. C. & Cole, R. D. (1916). Experimental investigations on the genus Drimys. Ann. Bot. 30: 359-368. Johnson, D. S. (1905). Seed development in the Piperales and its bearing on the relationships of the order. John Hopkins Univ. Circ. no. 178. Baltimore. Jussieu, A. L. de (1808). Sur les caracteres de familles tires des graines et confirmes ou rectifies par les observations de Gaertner, et specialement sur les Caprifoliacees et la famille nouvelle des Loranthbes. Ann. Mus. Hist. Nat. Paris 12: 285-303 (299). Leroy, J.-F. (1981). An unrecognized ancestral dicotyledon with a strobiloid flower is living today: Hedyosmum. XIII Int. Bot. Congr. Abstracts: 136. Sydney. -- (1983a). Interpretation nouvelle des appareils sexuels chez les Chlo- ranthacee's (Chloranthales, Magnoliidees). C. R. Acad. Sc. Paris, 296: 747-752. (1983b). The origin of Angiosperms: an unrecognised ancestral dicotyledon, Hedyosmum (Chloranthales), with a strobiloid flower is living today. Taxon 32: 169-175. Lindley, J. (1821). Collectanea Botanica. London. (1842). The Vegetable Kingdom. London. Meeuse, A. D. J. (1962). The multiple origin of the Angiosperms. Adv. Fron- tiers P1. Sci. 1: 105-127. (1963a). From ovule to ovary: a contribution to the phylogeny of the megasporangium. Acta Biotheoretica 16: 127-182. (1963b). Stachyospory, phyllospory and morphogenesis. Adv. Frontiers P1. Sci. 7: 115-156. Melville, R. (1963). A new theory of the Angiosperm flower: II The Androecium. Kew Bull. 17: 1-63. Patel, R. N. (1975). Wood anatomy of the Dicotyledons indigenous to New Zealand. 10. Chloranthaceae. 13: 141-148. Smith, A. C. (1972?). An appraisal of the orders and families of primitive extant Angiosperms. J. Indian Bot. Soc. Golden Jubilee Volume 50A: 215-226 (and roneoed sheet dated 1973 attached to reprints). (1976). Studies of Pacific island plants, XXXIII. The genus Ascarina (Chloranthaceae) in the Southern Pacific. J. Arn. Arb. 57: 405-425 (405). (1981). Flora Vitiensis Nova 2 (97-101). Sprengel, K. (1817). Anleitung zur Kentniss der Gewaichse. ed. 2 vol. 3 [2:2]. Halle. Swamy, B. G. L. (1953). The morphology and relationships of the Chlo- ranthaceae. J. Arn. Arb. 34: 375-408. 224 KEW BULLETIN VOL. 40(1)

-- (1955). Sarcandra irvingbaileyi, a new species of vesselless dicotyledon from South India. Proc. nat. Inst. Sci. Ind. 19: 301-306. -& Bailey, I. W. (1950). Sarcandra, a vesselless genus of the Chlo- ranthaceae. J. Arn. Arb. 31: 117-129. Takhtajan, A. L. (1969). Flowering plants: origin and dispersal. Translated by C. Jeffrey. Edinburgh. Thierry, R. (1912). Contribution ai l'&tude anatomique des Chloranthacees. Thesis. Evreux. Tieghem, P. E. L. van (1884). Traite de botanique. Paris. Vijayaraghavan, M. R. (1964). Morphology and embryology of a vesselless dicotyledon-Sarcandra irvingbaileyi Swamy, and systematic position of the Chloranthaceae. Phytomorphology 14: 429-441. Xi, Yi-zhen. (1980). Studies of pollen morphology and its systematic position in the order Piperales. Acta Bot. Sin. 22: 323-329 (Chinese with English Abstract). Yoshida, 0. (1957). Embryologische Studien fiber die Ordnung Piperales. I. Embryologie von Chloranthus japonicus Sieb. Jour. Coll. Arts and Sci., Chiba Univ. 2: 172-178. (1959a). Embryologische Studien uber die Ordnung Piperales. II. Embryologie von Chloranthus serratus Roem. & Schult. Jour. Coll. Arts and Sci., Chiba Univ. 2: 295--303. --(1959b). Embryologische Studien iiber die Ordnung Piperales. III. Embryologie von Sarcandra glabra Nakai. Jour. Coll. Arts and Sci., Chiba Univ. Nat. Sci. Ser. 3: 55-60. Young, D. A. (1981). Are the Angiosperms primitively vesselless? Syst. Bot. 6: 313-330. Cumulated Contents to Kew Bulletin Vol 40 Issued with Kew Bulletin Vol 41 part I (ISBN 0 11 242731 6) CORRECTION Page V the first eight lines should read Page Verdcourt, B. A synopsis of the Moringaceae...... 1-23 A new Cyphostemma(Vitaceae) from Kenya ...... 135-138 An introduced Sesuvium(Aizoaceae) in Arabia ...... 208 - Notes on Malesian Chloranthaceae...... 213-224 A new combination in Hallea (Rubiaceae- Cinchoneae)...... 508 - A new species of Foetidia(Lecythidaceae) from Africa ...... 635-636 A new species of grassland Psychotria(Rubiaceae) from Africa. 647-648 - New taxa of Rytigynia(Rubiaceae - Vanguerieae)...... 656

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