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BLUMEA 43 165-182 (1998) The myrmecophilous species Guinea de Wilde W.J.J.O. Rijksherbarium/Hortus Botanicus, P. of Myristica from New (Myristicaceae) O. Box 9514, 2300 RA Leiden, The Netherlands Summary The phenomenon of myrmecosymbiosis Guinea, and the new reviewed here. of their status A as a new words: key variety Myristica, in M. a New in Myristica, only occurring myrmecophilous to is myrmecophily species (M. dasycarpa, scribed Key is discussed. briefly sarcantha, M. Three verruculosa), some species myrmecophytes and M. liable to be confused with species in is (and resembling species) taxa subcordata New presented, described are var. from as rimosa is de- myrmecophyte. a ants, coccids. Guinea, myrmecophily, Introduction In the elaborate taxonomic species M. subalulata Miq. collections of this species and this character, together base petiole to is the almost petiole base, of the treatment only always 66, 384). Sinclair accepted M. subalulata in (I.e.: barium material of this common species has a wide for a Herbarium myrmecophytic. ant-swellings wing-like ridges major characters Sinclair (1968) the Myristica by show characteristic with the marked serve as genus identified as one defining wide sides at two twigs, from running his series Subalulatae and hence the her- sense, ecological in the range and shows a large variability. In my enumeration of was Myristica in New Guinea partly covered, for various morphological ties, viz. var. subalulata. var. larger fruit, as var. with the compared study acteristically with type-variety), since Sinclair's revision, ant-swellings 1995; and the present paper). twigs, ants and (i.e., character a generally distinct hollow on (De Wilde, species more the stem it is used in of New Guinea. Myristica bialata, M. mountainous plants), and now a twigs, have the conviction that an Myristica and in the , early fork, leading Including the three verruculosa. species, new to by to the species me also char- (De Wilde, ant-swellings to ant-holes), is in the be inhabited by a preponderant New Guinean species myristicaceous ant-plant described here, these dasycarpa, M.fasciculata, M. fissiflora, M. subalulata, and M. pedunculata (with potential key var. some- material has become few additional new and/or the presence of in var. more have been described the inhabitation or the visible species-level 1995) I in the variability several varie- paucifructa (infmctescences with few, peduncled inflorescences). Besides, because considerably available for this by accepting reasons, hagensis (coriaceous-leaved leptantha (with slender male flowers), what (De Wilde, 1995) M. ingrata, are: M. sarcantha, BLUMEA 166 The habit of housing with twigs the Kai and Aru ing paratively tendency and Plants of these ridged or species especially when the collected have species than 20 more twig portions some angled and of the inhabitedby ants, and when the mountainous M. subalulata cies, supposedly obligatory inhabited by hence have in M. bialata and in the twigs (i.e. As said above, still other inhabited never M. hollrungii, twigs or subcordata). Finally, look listed and entered in the With the AND scribed key STATUS of the treatment elements (1968) (with OF conceived widely now in two narrow wings running throughout the with circular in being an as result of the often swollen cies of run although some of the mostly have specimens just not MYRMECOSYMBIOSIS always ants be or Sinclair Miq. by leafbase, special one on de- two notes lines or as habit that twigs are are wings the to in the swollen are illustrated, species (I.e.: inhabited by to petiole base the do show them. It apical parts and seem to seems of the (also that the ants which two ex- lateral found in certain spe- myrmecophilous parts where they form portions twigs diagnos- hollow and are be absent from the seem to twig. Myristica hollrungii and the swollen on ants from the older, solid, striate parts. The base 393- family modifications that have been well-known and useful swellings often each side, the lines present modified into morphological very distinct in the present." is twigs non-myrmecophilous parts twigs and M. sulcata also have the are not inhabited by ants," be fewer in that such (such) portions and that the prefer them are they and mountainplants, without them portions showing been collected. On the other hand it may be that by M. inutilis, for convenience sake. OF ant-holes. From the petiole ant-swellings the younger or vaded yet cannot the wings M. lined solid and these have been smooth and also solid, becoming deformed. The each side of the on lines but lack the "... are not apparently are hardly (I.e.: 387) twigs with swollen parts of the well down from Horsfieldia) wings as but to myrmecophilous variously or do hollow subalulata is distinct from all other members of the portions apical portions lines which thin Such 64 and the ant-plant a twig fig. elongated Myristica tic characters. treme as " 396), I quote: evolved In his well as from leafbase of the length parts." (I.e.: 386). not or twigs), in such parts, the older apical parts quadrate, the ants Other spe- M. sulcata, the condition of the segregated species) sup- with hollow, swollen, ant-inhabited deformities, reddish- brown, smooth and often shining generally solid,..., but M. subalulata species are though myrmecophilous species (e.g. PHENOMENON THE ascertain and M. verruculosa). twigs with terete, myrmecophilous species to "myrmecophilous as like swollen ant-species (i.e., of M. subcordata with hollow or solid specimens HISTORY deceptively not species some to hagensis). dasycarpa hollow (partly) are ridged) twigs, short always present) not a con- actually sheltering var. ant-holes (and described M. may have species and hence these ants vegetatively may partly, by newly ants normal-looking, be can myrmecophilous species only they produce swellings (e.g. normally produce swellings, too com- twigs (with species lined (or or inadequate are the long, these (includ- and leaves of twigs in conspicuously more cm low-winged. However, or if the character is present. Moreover, posedly merely facultatively and twigs is exclusive for New Guinea without ant-inhabitationbut with fused with 1998 1, in hollow Myristica, perforation holes, Islands). lined be) No. 43, habit, with leaves generally stout to in ants ant-swellings —Vol. have not are them been in- absent in the district. Ants W. J.J.O. de Wilde: Beccari (1884) mecophila Becc., a of transcript servations cited is a published on of portion Bower (1889) publication by a and reproduced the slit-like holes, Clerodendron fistulosum, of which two each internode. Beccari is of a new aptation this situated, opinion access either side, one on of the protecting also Maschwitz et the spontaneously, i.e., al. (1994) report opportunistic discovered ant of case being as a as for M. 65 time and offer long one stomata and inhabited by are and notes myrmecosymbiosis in the in New Guinea found that the provide references hundred years ago, Schumann already published Myristica short essay a Myristicaceae, (including by ants the a notion pith several and subalulata assumed that the concerned to twigs, even plant itself The or occurring (1890, had 1897) of what is in some In essence tissue of the frequent galls, be resolved which are as myrmecophila well is called Warburg and growing twigs were even of of species internodes of the occurrence was not the way the were a twigs. fairly large always present the the (= M. opinion case, by were him in sterile definitely that in and that the ants ants disperse too to to on to are to discussed irregular is inherent preceded plants, be by the in fer- as is truly genetically a He Miq.). well ant-swellings ant-swellings the young big as located should have Myristica myrmecodomatia. Queries in the adult stage sometimes subalulata ant-species, particularly belong found stimulus from the of the true Becc. of whether the formation of question rather than are as in very young stages, but rather that and he myrmecosymbiosis of and M. phenomenon, apically twigs. ply discussed, sort Miq.) Myristica species. Swellings the on the (Capparis has Myristica species, viz. M. heterophylla K. Schum. (= M. hollrungii Warb., p.p. two and ?M. sites a opportunistic ant-spe- Warburg coccids in the ant-hollows, and further he assumed that these tile number related literature. pre-indicated he recorded the a buwaldae Jacobs, with simi- and Aru Islands). not already The sponta- al. (1996) describe et phenomenon only as from the morphologically on to (1889) on the Kai ant-swellings originate formed irregularly, Besides the habitation in plant." unspecific an space for nesting Maschwitz species. Likewise, Both articles species). About the by myrmecophila. cies; it is the first and apparently only myrmecophyte in Capparaceae some ad- an Clerodendron fistulosum on develop by themselves, well as myrmecophyte, Capparis a formed larly spontaneously holes species formed holes remain open for neously of different newly entrance Beccari in 1884 for this by as The form of the orifice in species. a new species, with extra-floral nectaries, of which the inflated hollow internodes myrmecophyte open the interior of to the upper end of Somewhat similar slit-like holes ants. Becc., grown in the greenhouse, without the presence of ants, held at that both the swollen form of the internodesand myrmecophila, publication a recent plants here also the internodes species, also would suggest that the initiative in their formationis taken case In requirement be found in M. to are the to are myr- Beccari's ob- myrmekophilous of the holes have become inherited characters of the origin the first 167 (I.e.: 396) notes concerning here: "Of the partly repeated swollen, and hollowed, and inhabited by ants, which gain are Guinea myrmecophilous plants, describing Myristica latersynonym of M. subalulata Miq. In Sinclair's a Beccari... is by New Myrmecophilous species of Myristica from regarded him as am- fixed as still a to and how the coccids, pass the ant-holes, enter the hollow nodes. Meanwhile I found in present inside the recent collections of M. ant-swellings. Notwithstanding subalulata the coccids the rather irregular frequently location of the BLUMEA 168 morphologically species, I well-defined ant-swellings myself regard the not or but little swollen cific character 1. partly two No. on species though and (see Fig. Myristica still x 0.5 subalulata ( Polak the 1998 twigs, always present less as a obligatory, in several good hollow twigs (and with ant-holes in the can serve as a taxo- ant-habita- bark) is useful additional spe- 1). Miq. present), ant-swellings, coccids, 1, apparently evident in part of the material, and hence quite Fig. 43, syndrome, following previous authors, nomic character. Besides, in other tion in Vol. MP var. and 750). subalulata. Male ant-openings; twig stem with immature partly opened to flowers (bracteoles show ant-cavities with W. J.J.O. de Wilde: About the biology of the coccids likely belong to KEY the genus and where experiments (without TO , This key is applicable only of the essentially more, leaves larger 20 c. also la. Plants the or more, with ant-holes Male perianth angular. study and more facts. greenhouse MYRISTICA with twigs im diameter mm low-winged or ridged, i.e. hollow twigs (perforations), frequently usually large, 10 mm c. swollen partly hollow, locally or winged, ± long (c. 7 or more not, or ridged, lined, or mm in M. not myrmecophilous, twigs terete, or lined, or solid angular, hollow, without ant-holes. Male perianth usually less than 10 with Twigs usually swellings Twigs (sub)terete, hollow, tion with ant-holes may 3a. and leaves Twig to 43 18 by (0.5—) 1 b. Plants ingra- without not Fruit cm 3.5 towards twig with long, stout, twig up to 8 apex (twig por- indumentum with hairs conspicuous mm dasycarpa diam. Fruit with shorter indumen4 perianth at anthesis in various stages of with certainty.) — split into lobes development; to halfway or bracteole small over; or flowers in large. (Fruit perianth split 5a. Mature male long only. 0.5-1 the in bud perianth long. mm 6a. Bracteole less, bracteole small, 2-3 oblong at or to or 4 cm according 1.5—2.5(—3) cm 2-10 mm the veins generally Aru Islands, not known not to long. 5 same to 0.2 with sterile apex (tubiform), lobes or subglobose, with ovoid, persistent glabrous Throughout Archipelago; or the ± 1.5-7 large, 1-1.5 synandrium inflorescence either of Lower leaf surface in the Bismarck anthesis mm 7 age. Fruit small — at long long, generally faint. subspecies) ellipsoid-oblong, or oblong-lanceolate Flower buds in the ing pedicel tertiary clus- mm anthesis for 1/6-1/8. Sterile apex of caducous. Fruit rather small, apex, ellipsoid or Synandrium obtuse, Fruit rather small, up of different stages acute or in bud ovoid perianth perianth splitting or for 1/3 into lobes for 1/3-1/4. b. Male one Lowlands of northern New Guinea 4. M. fissiflora (2 b. Male split 3 .. diam., leaf blade mm 2. M. less 11 ridged 9 10 c. long generally or absent or narrowly have been collected) stout, c. but long mm usually winged ant-holes present swellings, exceedingly cm. mm and with ant-holes, or tum 4a. Male ter or 2 b. Plants b. or and with ant-swellings, not, outside occasionally 5 c. without or ta)I 2a. the or not. myrmecophilous, usually OF i.e., twigs towards apex without ant-holes, inside hollow or angled, lined, SPECIES yield of species Warburg (1890) to from New Guinea (incl. Kai and Aru Islands), specimens to Field Targ. with ants) may the 169 resembling non-myrmecophilous species) size class, long cm Myzolecanium possible Guinea swellings, the Myristica i.e., MYRMECOPHILOUS (including or in syndrome concerned, and the coccids, little is known yet. According ants New Myrmecophilous species of Myristica from cm or mm, acute, same long ellipsoid, stage .. 6 with short tomentum, fruit- or minutely pubescent, New Guinea, incl. the Kai and 0-2100 7. M. subalulata (5 m altitude varieties, excl. var. hagensis) 170 BLUMEA b. Bracteole rather distinct. or No. 43, in anthesis. Fruit 5-7 persistent veins faint —Vol. 1998 1, long, (late) glabrescent. Tertiary cm Lowland, Papua New Guinea: Bismarck Archi- — 1. M. bialata (2 varieties) pelago 7a. Flower buds before anthesis in cording to age. Bracteole ovoid-ellipsoid, c. 2.5 inflorescence one persistent. long, cm of ginal arching and nerves may be confused with M. ern narrowly apex 3-10 mm apiculum); fruiting pedicel stoutish, veins tertiary subalulata.) long. opment). one perianth low the Irian inflorescence all of about the veins tertiary stout, same long. — mar- areas of north- ingrata (2 subspecies) size (same of develstage 7-8 interarching mm; and fruit not of 8 bracteole known.) situated well be- scar — Lowland of northern Jaya (West New Guinea) 6. M. sarcantha pedicel longer with bracteole Female flower pedicel perianth. minute distinct generally pedicel carnose, (with (Fruiting specimens Montane and lowland — 5. M. perianth. (Female flowers b. Perianth smaller, and faint. Bracteole caducous. Lower leaf surface with lines of lateral nerves and 8a. Male rounded ac- Fruit mm. Lower leaf surface with generally and southern New Guinea b. Flower buds in pedicel 2—5(—6) broadly or of different size generally Female flower c. 10 mm at scar the transition of Lowland, Papua New Guinea; Upper Sepik River pedicel 10-15 long; fruiting pedicel mm (W Sepik Prov.) area 3. M. fasciculata 9a. Plants montane, found b. Plants from lowland 10a. Male anth perianth b. Male perianth — 1 la. to about W Irian inside Jaya: 5(-6) c. 2 cm lobes hardly finely warty; 6. Lower leaf surface or Archipelago 1.5 c. var. cm — and Manus Island Papua — 1. M. bialata (2 ... densely finely warty; lobes splitting peri- altitude m not known) less than 10 (M. subcordata long, dry pericarp Dayman, c. Papua 20 Milne long, dry pericarp 2(-3) — 8. M. verruculosa mm — Lowland gigacarpa mountainous) var. mm thick. Bay Prov. (Flowers New Guinea: Western not 11. M. ... thick. Male mm 12 long. Twigs distinctly angular. Twigs ± terete, lined 14a. Leaves below — or not. — (sub)glabrous (i. 10-14 Highlands Prov glabrescent. — Mostly glabrescent, at first with weak Fruit 2.5-5 cm long, mm or (sub)persistent and medium altitudes (but see hagensis tomentum. M. subcordata var. anthers 8- subsessile, largely late- lowland marshy forest; 0-1000 dense long, dense minute long, m altitude 9. M. b. Leaves below with mm 14 montane tomentum). Mature male perianth ellipsoid-ovoid, 4-6.5 10, bracteole subpersistent. — 14. M. sulcata Lowland or e., or 13 pachycarpidia perianth Lowland . known.) 12. M. subalulata var. b. New varieties) Montane, 1500-2000 m altitude some taxa bracteole caducous. 13a. hagensis splitting peri- mm, glabrescent. Bird's Head & Bomberai Peninsula New Guinea: Mt Papua b. Fruit 12. M. subalulata . . 1/3; anthers 6-9. Lower leaf surface with persistent dense tomentum. montane, 0-1500 E altitude inside towards the base perianth (in 12a. Fruit m 10 not or Twig strongly angular. b. Male 2000 area about 1/8; anthers 5 to Guinea: Bismarck anth at c. — Dry gigacarpa) hollrungii land forest of low 15 W.J.J.O. de Wilde: Myrmecophilous species of Myristica from 15a. Leaves of medium size class, 3-4.5 lipsoid-oblong, Leaves medium or 12-24 large, mm anthers long; 1-3 long; fruiting pedicel cm b. mm mm 12-40 (male flowers known in M. subcordata not mm long; fruiting pedicel Mature male anthers 5 long; var. 1-10 perianth 171 in bud el- Fruit 2.5-5 subsp. perianth papuana in bud narrow, 6; bracteole caducous or gigacarpa mm Guinea persistent. 10. M. inutilis long. (6.5-)7-8(-10) cm bracteole long cm almost tubiform, (3.5—)4—5.5 Mature male long. (3—)5; New and rimosa). Fruit var. long 13. M. subcordata (4 varieties) ENUMERATION 1. bialata Warb., Myristica Notes This — Lowland — The species are may be due to is (1891) 308; W.J. de Wilde, Blumea 40 Arbor 0.5-1 pilis 10 m c. cm by facultative W. J. de Wilde, habitu longae 19 cm spec. fructus in latae, brown, with scattered mm ± mm high running concolorous Miq. with stems cases their ab- however. nov. — valido, ramunculi apicem stout, 8-10 high. Twigs spicuous wings 2(-3) because hollow myrmecophyte, sicco longis. —Typus: Regalado & mm Archipelago. Sinclair (1968) in M. subalulata inadequate collecting, myrmecophila ad 47 a of Manus Island and Bismarck included manifest in part of the material; in these latter not Myristica dasycarpa nae area was species possibly ant-perforations Tree, 13 W.J. de Wilde, 1.c.: 264, f. 2b. brevipila var. Distribution 2. Jahrb. bialata W.J. de Wilde, 1.c.: 264, f. 1, 2a. a. var. sence Bot. TAXA f. 1,2a, b. (1995) 264, b. OF MYRMECOPHILOUS 2 Fig. 10 versus c. 3.5-4 Takeuchi mm longi cm 1520 diam. alati, indumento (L), Papua New lami- velutino Guinea. diam. towards the apex, subterete, with at both sides from petiole lenticels, twig partially to con- petiole, bright somewhat swollen and hollow, ant-holes present; bark of twigs lower down (where bearing the fruits, below the leaves, stem 15-20 mm by 13-19 cm, cracked and diam.) coarsely rectangular brown. Leaves: blade membranous broadest at or thinly chartaceous, ± about the middle, base densely cinnamon per side, tomentose with interwoven hairs 0.1 dark dots absent; midrib slender, flat or at an angle (—0.3) slightly grey- dry, oblong, (narrowly) cordate, broadly acute-acuminate; upper surface drying olivaceous(-brown), papillose; flaking, brittle when mm; not or raised above, of 70-80° with the midrib, flat, lines of apex lower surface indistinctly nerves 20-25 interarching fairly dis- tinct, tertiary veining finely trabeculate, indistinct; petiole comparatively short, thick, almost pulvinate, rescences (known brachyblasts c. subsessile, 2 or fruit 5 15-20 by 8 from the mm glabrous; diam. Male 3 per sessile ovoid-ellipsoid, rufous-rusty mm, infructescences) base terminal leaf bud not seen. of the Knemai.e. type, inflorescences knotty infructescence, not on known (but the older broadly rounded, apex narrowly velutinous indumentum with hairs 0.5-1 note 2). Fruit below the leaves; rounded or mm; inflo- wart-like see twigs Female woody (sub)acute, dry pericarp with strongly BLUMEA 172 Fig. 2. Myristica dasycarpa tomentum of lower leaf cences, x 0.5 Vol. W.J. de Wilde, surface, x 20; c. a. lower 43, Habit No. of portion (Regalado& Takeuchi 1520, type). 1, 1998 portion of older of leafy twig, twig x 0.5; b. with ramiflorous detail of infructes- W.J.J.O. de Wilde: creased much shrunken (and apparently stoutish, 2(-3) mm long; New Myrmecophilous species of Myristica from seed mature drying), on oblong, 2.5 c. 3 c. mm thick; 173 fruiting pedicel the aril long, greyish, cm Guinea impressed into the seed. Field-notes Distribution Papua Tree — fruits rich brown Known — New Guinea, East around Waskuk, at Notes Ecology 50-70 villages, is species see Ambunti the subsessile fruit, tree; on side a 2. Male flowers of this forest 850 at long, cm 1 to c. species m, mm described briefly long, pubescent, of various 2 c. ing mm, on the The to M. 1.5 c. Brass Idenburg River, in flood here. In L this collection consists of a portion separately added subalulata. The immature male flowers follows: inflorescence bracts specimen mm, with a a wart-like 0.5 long, almost Twig blade to 1 c. Plants published and leaves 43 18 by mm 1 mm, can ± mm, acute, anthers be inserted in the in Blumea 40 exceedingly cm. Fruit less c. 6 mm 6 flower (buds) mm, bracteole 14 woody (not collaps- (?); androphore c. 3.5 (1995) stout, cm general key 241 twig long, to New Guinean after lead 22 — narrow, towards apex with c. Papua 10 mm diam., leaf conspicuous indumentum with dasycarpa Fruit with shorter indumentum de Wilde, Blumea 40 New species follows: as M. stout. fasciculata W.J. Distribution by by long generally Myristica 3 3.5 or glabrous. Myristica dasycarpa as c. 5-6 3-4 brachyblast, subumbel of 5 thick, splitting bud for about 1/3 (?); androecium c. long, synandrium hairs certainty. on mm Myristica 3. as conspicuous valves of b. belonging glabrescent, drying), a. most ellipsoid, 1 22-bis to bud c. 3. the trail Garuka with for the group the stages of development; male flowers: pedicel slender, 4 mm area apical; (immature) c. mm Hills, long. known with not are possibly belongs fruit, the latter obviously be from ridge with inflorescences with immaturemale flowers and leafy twig can Waskuk habit, with thick alate twigs and large leaves, 13706, from Irian Jaya, 4 km S W of Bernhard Camp, plain 2), collected in northern note subprovince, distinct among the New Guinean myrmeco- stout 3.5-4 indumentum: velvety with hairs of a of nutmeg. 4° 11' S, 142° 44' E. readily because of its philous Myristicas by from the type (but only beneath; tomentose strongly smelling altitude; fruits July, 1995. m This 1. — crushed dried fruit Subcanopy — leaves brown myrmecophilous, m, Sepik Province, and Garuka Langu Habitat & and 10 c. orange-brown; to (1995) Guinea, Upper Sepik 23 278. River area; in forest at low altitudes. Note — The limited number of collections known suggests that this obligate myrmecophyte. 4. Myristica fissiflora W. J. de Wilde, Blumea 40 ( 1 a. subsp. fissiflora; b. subsp. W.J. de Wilde, 1.c.: 282. kostermansii W.J. de Wilde, 1.c.: 283. 995) 28 1 . species is an BLUMEA 174 Fig. 3. Myristica —Vol. sarcantha W.J. de Wilde. a. 43, Habit of perianth lengthwise opened, bracteole caducous, x 6 No. 1, 1998 portion of ( McDonald leafy twig, & Ismail x 0.5; b. male 3800, type). flower W.J. JO. de Wilde: Distribution Northern parts of Irian — forest, 0-400(-1000) Note was — This Myrmecophilous species of Myristica from a. subsp. ingrata; b. subsp. Distribution Notes New Guinea; in lowland Papua W.J. Part of the material obligate myrmecophyte. an Sinclair (1968). Miq. by de Wilde, Blumea 40 f. 2c. (1995) 290, W.J. de Wilde, 1.c.: 291. velata W.J. de Wilde, 1.c.: 292. limestone, in the is species apparently Myristica ingrata 175 m. included in M. subalulata 5. and Jaya Guinea New — — Most of New Guinea, in lowland and lower 500-800 m at This is with hollow, obligate myrmecophyte, an Occasional twigs. forest; also montane on (subsp. velata). conspicuous irregularly shaped usually swollen portions look like apical swellings insect-galls. The rather small male perianths (6.5-7.5 mm long) reminiscent of M. are hollrungii Warb. 6. Myristica sarcantha W.J. de Wilde, spec. Ramuli cavi formicis incolentes. longo (4-)5-6 mm 3800 (L holo; Treelet, 4 m GH Irian but ant-holes (hardly striate), petiole subglabrous, minute scattered whitish flat above, nerves interarching ole rather long, pale, by 3 ly minutely mm, Male flowers stout; (not seen), its scar oblong-fusiform, tapered, apical 1/3 glabrous at an upper mm angle i.e. apex surface acute, sterile apex 0.5-1 mm c. in the stage of with stellate hairs mm, mm, 8 by 2.5(-3) not seen. cuneate, apex and with extremely at both surfaces; by 3-angled 3 to 5 not seen. mm development, (less than) mm, pedicel; petiMale diam., the extreme- 0.1-0.2 mm. bracteole caducous mature perianth in broadest above the middle, acute, base 0.6-0.8 mm, mm with lines of lobe-sutures well- thick; androecium stout, 12-13 anthers 12 long; androphore but with very minute (less than 0.1 Female flowers and fruit smooth drying green-olivaceous, venation distinct stout, 7-8.5 5(-8) seen), not brown, glabrous; terminal leaf bud at suture subterete of 70-80° with the midrib, flat, lines of slightly differing long, apex, hollow (see field- brown dots absent; midrib slender, almost tertiary by (4-)5-6 long, synandrium oblong, contiguous, twig about 1/3-1/2 from the apex of the portion, marked, lobes 4-4.5 tip; mealy pubescent 15-16 mm Leaves: blades membranous, minutely irregularly papillose pedicel short, at but of leaves, sessile, wart-like, scar-covered, scattered ± 15-16 McDonald & Ismail broadest above the middle, base slender but distinct; 25-30 Typus: diam. towards the mm inconspicuous. appressed hairs, flowers in subumbels of 2-5, mm cm, 20-22 per side, nerves inflorescences axillary bud stout, 5-6 rounded with acute-acuminate lower surface — petiole, bright brown, partially to 8-15 by 3 grandes carnosi, perianthio glabrous (glabrescent, seen, 22-32 Fig. Jaya. lenticels small, scattered, obovate-oblong, broadly not — apicaliter triangulari. high. Twigs moderately with shallow lines from notes) Flores masculi lato in alabastro iso), nov. c. 4 by or 13 (i.e. 24-26 thecae), 1.5-2 mm, appearing as mm) pale stellate hairs in lower 2/3. 176 BLUMEA Field-notes Small — Vol. tree, 4 understorey No. 43, m 1998 1, tall, branching above and below, lateral branches whorled; trunk straight, cylindrical, with muricate bodies that attract 3 diam.; lateral branches covered cm ants; flower tubular, tube aggressive and pale yellow green, limb segments reflexed. Distribution from the type collection, only Known — at Sarmi, northern Irian Jaya (West New Guinea). Mixed primary lowland forest, canopy 25-45 1-3 km N of Sewan plain, on the Waske River, 2° 4' S, Coastal high. m 138° 46' E. Altitude 10-20 m. McDonald & Ismail 3800, male flowers June 1993. Notes ered 1. Apparently belonging — alulata and readily its by to the group robust large, is M. verruculosa, with flowers species nomically distinct well as as in general of ant-inhabitedspecies of M. subflowers. Another stout-flow- carnose larger mainly by habit M. sarcantha is close a M. to longer pedicel. Taxo- fissiflora (with deeper lobed, less thick-fleshy male perianth, and with the bracteole strictly apically and also in pedicel), general habit close to M. fasciculata (mature the at male flowers not known, but immature flowers much different). 2. sarcantha Myristica be inserted in the can (De Wilde, 1995: 242) by replacing lead 26b 26a. as general key to New Guinean M. (unchanged) b. Flower buds in 26-bis nerves Male a. and veins tertiary perianth below scar inflorescence all of about the same stout, interarching distinct generally carnose, 15-16 mm perianth. (Female flowers and fruit 7-8 long, pedicel known.) not — mm, bracteole Lowland, N Irian M. sarcantha Perianth smaller, pedicel longer Female flower perianth. pedicel with bracteole c. 10 mm at the transition of subalulata Myristica Miq., Ann. Mus. Bot. Lugd.-Bat. 2 . .. pedicel 10-15 long; fruiting pedicel Lowland, Papua New Guinea, Upper Sepik River (W Sepik) 7. of 26-bis Jaya b. ingrata size (same of develstage Bracteole caducous. Lower leaf surface with lines of opment). lateral one species follows: and mm. — M. fasciculata (1865) 47; W. J. de Wilde, Blumea 40 (1995) 330. With five varieties, of which var. hagensis W. J. de Wilde is considered as not myr- mecophilous. Distribution tude 0-2100 Note — var. alate ex Miq., — 1. regarded as a (I.e.: 274) a true, obligate myrmecophyte, non-myrmecophilous species). perforated hollow ant-swellings Al- in the twigs. M. costata myrmecophila Myristica Blumea 40 (1995) 274 me below in the list of subalulata; W. J. de Wilde, 1.c.: 330, f. 2d. Notes Kai and Aru Islands; alti- including subalulata Miq. is considered as hagensis (see Synonyms: ?Myristica Zipp. in New Guinea, all herbarium collections possess (shallowly) a. var. Wide-spread Myristica except for most — m. - costata Fig. 1 Becc. Warb. [Bot. Jahrb. 18 (1893) 191; W.J. de Wilde, type Hellwig 247, Bt from Sattelberg] is doubtful; it synonym of M. subalulata Miq. this is — Warb., ?M. heterophylla K. Schum., M. macrophylla unlikely. by Sinclair (1968), but as was explained by Myrmecophilous species of Myristica from W.J.J.O. de Wilde: 2. K. Schum. Myristica heterophylla land (1889) 45] is doubtful a name. K. Schum. & [in Hollrung, The type material consists of Guinea New 177 Fl. Kaiser Wilhelms- specifically different elements, but the lectotype, Hollrung 648 (male fl.), designated by Warburg (1897: 489, 11) is lost in B and I have t. Sinclair description, swelling 11, however, depict t. plant a given vegetative subcordata Blume 1880, the latter belonging 3. When MP 750 8. variety imagine homonym of M. coccids found within the sedentary Fig. Irian m lines from without was hidden, the baby regarded at present accept de Wilde. The name opened ant-cavity some of Polak 20 minute young deemed to be able creatures taxon W.J. de Wilde, 1.c.: 331. bialatae masculi affinis, sed foliorum interiore verruculoso nov. Fig. inferiore pagina differt. — — Typus: collected). 4 tomento Polak MP persistenti denso, 1283 (L holo; BO Jaya. high. Twigs petiole to than 0.1 base stout, 4-5 mm diam. towards the apex, subterete with faint petiole, light brown, partially hollow with incidental ant-holes, mm, at first with minute brown bright tomentum or oblong, broadest at or broadly rounded, sometimes shallowly nerves the midrib, line of 20-40 mm, by 5 mm, mm or less, not above the middle, 20-32 cordate, apex papillose, acute or glabrescent, nerves brown to 5 mm, as glabrescent; at an angle by acute- dense slightly of 60-80° with faint; tertiary veining indistinct; petiole stout, the twigs; mm or low the leaves; of the Knema- type, i.e. in male peduncle without dots; midrib flat or 20-23 per side, flat or sunken above, interarching with dense brown stellate hairs 0.1 a stout of older bark striate with scattered lenticels. Leaves: blades of pale hairs 0.1 raised above, to 12). acuminate; the upper surface olivaceous, the lower surface with persistent tomentum M. heterophylla Fern.-Vill., non-myrmecophilous (see below, as chartaceous, elliptic-oblong cm, plant but the above with this combina- de Wilde, 1.c.: 330. swellings, early glabrescent, hairs less hollrungii, 1. verruculosa W.J. de Wilde, spec. Myristicae perianthii an ant- Knema. W. J. de Wilde, 1.c.: 332, f. 2e (once Tree, 15 a W.J. pedunculata iso), M. to and well-developed leaves; normal appearance in what I W. J. de Wilde, 1.c.: 332. Myristica 8-13 to paucifructa var. e. var. also W.J. is leptantha c. var. to belong cannot invalid later with distinct feet see hagensis var. This d. of the an are a well removed, it appeared that underneath a flock of around; move b. one was provided ones with hollow, non-alate stem, and morindiifolia (Blume) var. K. Schum. is heterophylla can characters do not, and I according hollrungii. Warburg's drawing with distinct ant-hole, and reduced leaves in between tion of traits. The reduced leaves M. which is in BO, duplicate is M. asserts the inflorescence and separate flower as the seen not and which he (1968: 410), bracts terminal leaf bud slender, less. Inflorescences simple or forked, to axillary 10 mm of 15-20 or long, be- with minute, caducous; male inflorescences 178 Fig. b. BLUMEA 4. Myristica leaf, x 0.25; bracteole drawn verruculosa W.J. de Wilde, c. male flowers, separately, x 6; the 1283, type; e, f: Polak MP 697). a. mature d. male flower, perianth lengthwise opened, MP —Vol. x 6; 43, 1, Habit of leafy perianth inflorescence f. No. portion of in bud with twig 1998 twig with male inflorescences, x 0.5; lengthwise opened, (sub)persistent one with flower blooming, infructescence, x x 0.5; 0.5 e. female (a—d: Polak W. J.J.O. de Wilde: with clusters low brown long as of the with hairs 10-11 perianth, acute, 4-4.5 oblong, (10—) 12 by acute, not acute, 4-5 glabrous, acutish, c. 4 bracteole inserted apical long, mm mm; the at i.e. 8—9(?) (i.e. c. mealy, ovoid(-oblong), mm bracteole oval, ing, 5 c. (-0.2) hairs 0.1-0.2 slightly hooded, (inch stigma mm. c. lobes 5 pedicel 2(-2.5) than 0.1 2.5 1.5 mm) mm by 3 tomentum long, with mm, mm, tomentum rusty, of hairs 1-3 Clear exudate. Leaves pale c. 0.5 mm; mm 0.8 by mm, Lower 1/3 of the visible with perianth lobes a of mm, taper- hairs 0.1 appressed 3.5-5 long, in bud 1.5 c. ovary ovoid, base broadly (ovoid-)oblong, seed at sub- sterile apex 5-6 mature 4-5 by to buds of various size; long, (sub)persistent, thick, fruiting pedicel mm mm. perianth mm 8 c. perianth fleshy, ± contiguous, finely irregularly warted, clearly apical portion, c. male 1/3, as rhomboid, long, synandrium oblong, of clusters of 1-3 fruits; fruits Infructescences rounded, apex acute, pericarp mm; or the apex bluntish the bud for 17 thecae), in lower half with hairs less narrowed in the stout, about mature tapered, androphore slender, cylindrical, surface inside conspicuously minutely yel- age, pedicel pedicel; splitting lens. Female flowers in sessile clusters of 5-10, with tomentum to 179 Guinea ovate-broad-triangular the base somewhat mm, anthers mm, 0.2-0.3 minutely pale pubescent perianth mm, according Maleflowers stout; mm. thick; androecium (8—) 10 mm 1.5-2 by 1.5 4-5 lobes 3-angular, 0.5(—0.8) by 0.1 c. (sub)persistent, mm, in bud sutures of different size 10(-20) flowers, pubescent New Myrmecophilous species of Myristica from 2.5 by ellipsoid-oblong, cm, dry 2.5-3 cm long. Field-notes 1 c. cm Distribution lrian Jaya (West the latter in the ISIR research forest on Notes Bombarai Peninsula and Bird's Head, Guinea): E of limestone area Sorong, verruculosa is related Myristica and M. subcordata, and hollrungii, species New area, of the Bismarck is distinct from all these perianth, however, Archipelago by distinct ant-holes in the var. areas, lar velata) equally persistent twigs, glabrescent and much structure and further within the differing by lower leaf surface; M. by having (sub)glabrous leaves; less (5 or Myristica new and M. or close bialata has M. to perianth. a a ingrata, M. bialata, new species A warty inner species much with species similarly is a species re- mostly (ex- of swampy M. subcordata, with simi- more narrow stout male perianths, male flowers, but dif- 6) anthers, shorter perianth lobes, and the (almost) absence a glabrescent lower leaf surface, densely short felty in species. 2. Because of the resemblance in habit of the present incomplete secondary much smaller flowers and hollrungii leaf indumentum, has much smaller and of a warty inner surface; it has the present m, resembling fissiflora (subsp. fissiflora), with smaller flowers and with caducous bracteole. fers 450 in East New Guinea. The present the fine warty is shared with M. to especially morphologically lated, but with deeper cleft male perianth. Myristica ingrata is cept at c. Pawiah ksiah. name l. , a yellow brown, soil. clay Vernacular M. brown below. Flowers long. sterile specimens species with other species, may have been mislaid especially underM. subcordata hollrungii. 3. This new species can be inserted into the key to New Guinean species of Myristica (De Wilde, 1995: 258) by changing leads 163 & 164 as follows: 180 BLUMEA 163a. 43, Vol. 1998 1, No. M. simulans (unchanged) b. Male perianth about (or 12 mm c. nearly) as long long less, with hairs 0.1-0.5 or the as long; androphore mm synandrium, mostly (partly) pubescent. various 163-bis a. Leaves of medium size, 3-4.5 ellipsoid-oblong, able in size, 2.5-5 mm i.e. 12-24 long; anthers persistent; long; fruiting pedicel cm Mature male long. cm bracteole 1-3 mm b. Leaves medium large, or 1-10 larger. Fruiting pedicel 164a. Mature male 12-35 in bud perianth mm cm b. Mature male perianth thers 8 oblong, (subsp. papuana) perianth in bud much 164 almost ± narrow, or tubiform, (6.5-)7-8(-10) mm 6. Perianth inside towards base smooth. 10-12 mm 9. Perianth inside towards base or Mature male long, fruiting pedicel ± in bud Fruit vari- long bracteole caducous; anthers 5 Fruit (3.5-)4-5.5 long. cm perianth (3 —)5. long M. inutilis long; Fruit 163-bis 1-10 mm M. subcordata long .. bracteole (sub)persistent; long; an- Fruit unknown finely warty. M. verruculosa Collections seen Polak MC 697 (female fl., fr.), — PolakNT 11450 (sterile ecological plot-specimen); de Polak MC 1283 (male, Vogel 9693 (fr.); Moll type), BW13007 (male fl., fr.). ENUMERATION (liable to 9. OF Myristica hollrungii Warb., Blumea 40 NON-MYRMECOPHILOUS be confused with Mon. TAXA myrmecophilous taxa) Myrist. (1897) 490, t. 19, 1-2; W.J. de Wilde, 289. (1995) albertsii Warb., M. Synonyms: Myristica K. Schum. heterophylla (p.p.), M. euryo- carpa Warb. Note but it ficially as — This species sometimes has hollow be inhabited never seems to it may resemble closely related, as some by twigs ants, and it and never forms of M. subalulata discussed by has Miq., Sinclair (1968:411). lined usually or ridged twigs, ant-perforations. with which it is Super- regarded However, it is quite distinct in its male flowers and fruit. 10. inutilis A. Myristica (1995) Gray subsp. This variable taxon can hardly confused with those of the very specimens never on papuana (Markgr.) W.J. de Wilde, Blumea 40 293. may recall hollow, nor be taken myrmecophytes ridged, and at as myrmecophilous, variable M. subcordata in vegetative but specimens (see below) can be of which certain habit. Branches of M. inutilis are present the species is somewhat arbitrarily separated male flower characters. 11. Myristica pachycarpidia A montane angled species only and hence subcordata var. superficially gigacarpa W.J. de Wilde, Blumea 40 (1995) known from the type. It has solid resemble certain also is 312. twigs which are strongly myrmecophilous.Myristicas. Myristica mountainous, but has subterete twigs. W.J.J.O. de Wilde: 12. Myristica This subalulata variety M. subalulata as Mt area Hagen Miq. Sinclair accepted by to create 13. coriaceous leaves. This variation was by Myristica subcordata Blume, by which the absence of discussed briefly collections from the some Highlands Prov.) up with other materialof M. subalulatabut differs and 181 differentiation in the variable some It consists of (1968). New Guinea, Western (Papua Guinea W.J. de Wilde. Blumea 40 (1995) 330. hagensis var. described in order was New Myrmecophilous species of Myristica from by link obviously ant inhabitation Sinclair (1968: 394). 1 (1837) 186; W.J. de Wilde, Blumea 40 Rumphia (1995)333. Distribution Synonyms: b. (notably var. morindiifolia). subcordata; W.J. de Wilde, 1.c.: 333. a. var. Warb. in New Guinea Wide-spread — Myristica fatua W.J. de Wilde, 1.c.: gigacarpa var. known from the type Only Houtt. subcordata (Blume) var. ?M. Miq., wallacea Warb. keyensis var. and 333 one additional collection in montane New Papua Guinea. c. var. morindiifolia(Blume) W.J. Synonyms: J. Sinclair, d. var. dense BO), Twigs Irian ridged nor rather flat and pale by or 5-9.5 contrasting Knema-type, c. 0.1 at paler mm the base ellipsoid, with the 2.5-3 Known Note — eral habit Ecology on — limestone Myristica as well aspera (L; only as and fissured iso c. about the middle; pale yellowish short, Female flower 1 mm at by first nu- oblong- or nerves brown dense 5(—10) by long. Fruit + 3-5 mm, sessile, ovoid-oblong, of hairs c. 0.5 2.5- solitary, ellipsoid, broadly rounded, top subacute, dry pericarp mm; c. 3 fruiting pedicel under the fruit, scar mm stout, of bracteole cm. from collections from Polak 697; Ridsdale CER 2342\ de Habitat & 9757 twigs. Inflorescences (female) of perianth persisting from forest or high; petiole colour of the with hairs base cm, long, frequently — conspicue Vogel rough, flaky 1-2 Distribution de diam. towards the apex, bark tomentum seed cortice Typus: coriaceous, elliptic-oblong, oblong, 2- (or 3-)flowered. sessile, 1.5-2.5 mm thick, with powdery (light) brown mm — conspicuously broadest cm, with the narrowed towards the apex, 3.5(-4) by juventis distincta. sunken above; lower surface with indumentum of hairs dark brown, aliis in ramulis pallenti lined, solid, 5-8 lenticels. Leaves: 17-30 pairs, not seen; morindiifolia (Blume) var. nov. verrucoso-fissurata soon elongated short-felty the fatua Bird's Head. Jaya, terete, not lanceolate, 25-35 var. subcordatae Blume varietatibus lenticellato blackish brown, merous M. M. multinervia A.C. Sm. rimosa W. J. de Wilde, Ab M. de Wilde, 1.c.: 334. Myristica morindiifolia Blume, Forest on bedrock; Vogel 9757', sandy 50-300 or m western Bird's Head, Irian Jaya: Schram BW 6099. clayey soil, presumably altitude. Fl. & fr. in subcordata is, like M. inutilis, accepted as in details of flowers and fruits, e.g. in the all collections May. very variable, in gen- length of the fruiting 182 BLUMEA and pedicel solid length of and is certain myrmecophytic species. and the present var. Of both varieties 1998 1, may in has species mostly completely devoid of ant-inhabitation.Stout completely morindiifolia, var. No. 43, of the hairs of the indumentum. The (sub)terete twigs, especially Vol. general appearance As remarked with the of description rimosa, their taxonomic status is provisional and sufficient material of male flowers is specimens, be confused with those of var. not gigacarpa, at all settled. Both varieties may lacking. represent separate species, resembling M. subcordata. Var. rimosa also may be vari- a ety of M. verruculosa. subcordata Myristica 1 mm 14. long) Sing. Habitat & sulcata Warb., ficially — distinct clearly the by rough, flaky apparently persistent perianth (with Monogr. Myrist. (1897), and lenti- hairs about — lowland dryland 334. 40(1995) forest. with rather stout, solid, species confused with In 19, f. 1-2; J. Sinclair, Gard. t. in New Guinea. Wide-spread Ecology This — The 23 (1968) 396; W.J. de Wilde, Blumea Distribution Note twigs. under the fruit is remarkable. Myristica Bull. rimosa is var. cellate-fissuredbark of the sharply 2-angled twigs may be super- myrmecophilous species. a ACKNOWLEDGEMENTS This article work of the northern ISIR Irian as the by project in discovery West Bird's of Myristica taxa not for the translating usual the fine Irian Head, Jaya through cooperation Veldkamp (L) vided incited was of the diagnoses of Bogor the seen before, Jaya (1994-1996) new and Harvard taxa collected and during field- general collecting Herbaria. into Latin. Mr. J.H. I thank Dr. Os (L) van in J.F. pro- illustrations. REFERENCES Beccari, O. 1884. Piante ed illustrata Bower, De da O. ospitatrici F.O. 1889. Proc. Phil. Wilde, W.J. J. O. ossa Beccari. Malesia Soc. piante formicarie 2. Instituto della Malesia Sordo-Muti, Genova: e della 37, Papuasia descritte 1. t. 18: 323. Glasgow 1995. Census of Myristica (Myristicaceae) in New Guinea anno 1994. Blumea 40: 237-344. Maschwitz, U„ K. Maschwitz, U., B. J. Dumpert, (Capparaceae), Jacobs Fiala Moog, a new & J.V. Lafrankie myrmecophyte K.E. Linsenmair. unspecific myrmecophyte & Azarae. I.Hj. from Borneo. 1994. Clerodendrum from Borneo with 1996. Capparis buwaldae Blumea 41: 223-230. fistulosum (Verbenaceae): spontaneously opening domatia. Blumea 39: an 143— 150. Schumann, Sinclair, J. Malesia. K. M. & M. Hollrung. 1889. Die Flora 1968. Florae Malesianae Gard. Bull. Warburg, O. 1890. Warburg, O. Sing. Beitrage 1897. Nova forscher68. 23: zur 385, Monographic der genus Myristica in 45. Berlin. Malesia and outside f. 64. Kenntnis Acta Abh. von Kaiser Wilhelmsland: Precursores XLII. The der papuanischen der Kaiserl. Myristicaceae: Flora. Leop.-Carol. 84-88. Engl. Bot. Deutschen Jahrb. 13: Academie 309, 310. der Natur-