36 The BLUMEA 53-61 (1991) tuberous epiphytes A of revision the of Institute, P.O. 4: Squamellaria M.H.P. Christensen Research Rubiaceae Jebb Box 305, Madang, Papua New Guinea Summary Squamellaria Becc. cies is reduced. Confined The the to is revised to include three genus on account stigma, of Hydnophytum Jack, fringed plates (squamellae) of the tuber has revealed Squamellaria species a has the holes the tuber wall, of S. thekii in ties. Each cavity in outline. crenate appears cavity is and the base of the a Girdle Chambers interconnect while along dorsally and their and basic encircling at gives rise rise apically arise only at '/3 of the tuber, are most stems are In apparently arising appears to to 'Deep-lying being between each of be associated with the formationof thickened and unbranched. with cylindrical, are similar cavi- recognised (Jebb, of This chamber entrance and edge, blind-ended and 'Superficial holes. The to Smooth Walled From its inner Smooth Walled Tunnels' stems to entrance a new in that the arise from the the apex holes cavity. one (fig. 3); In this (fig. 1). stems apical '/ 2 (fig. 2). The from younger densely waited, previous cavity. stems are more remote ring The Hydnophytum. it is Hydnophytinae clustered close S. major the capitate structure, which fur- (fig. 1). their basal numerous the tuber S. imberbis the apex. In slender and branched. stem rise origin- closely related, essentially ring to a along also differs from the rest of the Squamellaria not genus structure: 'Girdle Chamber' to is the species, the tuber. Between these rings of Chambers' which overlie the Girdle Chamber of the gives spe- swells outwards, such that the tuber apical edge. Ventrally they wall the Girdle Chamber existing the tuber apex, between the last formed older cavities basally they give three most and Several distinct chamber types to one bony endosperm. tuber contains several, superficial planar all but encircles the tuber, and cavities same particular, A mature stem. just which it is to embryos produced sequentially 1985). Each cavity has with remarkably complex cavity rimmed holes, regularly spaced, of which new, and inside the corolla tube, the the taxonomic 'moat' between this strengthens tuber of all rings this genus from of the four one & Jebb, 1991). Beccari (1886) Hydnophytinae (Huxley and the curved pyrenes with short Study ther species, Fiji. Fijian Islands, Squamellaria, eastern smallest genus of the ally separated is endemic to to stems are another, thus each species do the one stem stems 54 BLUMEA 1. Schematic Fig. Becc. a. cavity; Tuber e. chamber; The f. h. deep lying stigma waited and anthers are of Scale bar opening or sile inflorescences and also Squamellaria, Squamellaria 10 stigma shows is and capitate Wise. finely fringed and the large capitate No observations of interesting that in the Fijian species of noc- solitary relatively large (110-163 |im) four-colpate pollen, occur of later presenter mechanism. The pollination. have been made. It is (A. Gray) smooth walled by Rosemary heterostyly, a imberbis d. entrance hole superficial g. Drawn cm. above the anthers, suggests production scent of preceding cavity; = exserted. The species 1991 1, girdle chamber; often curved corolla tube suggests moth turnal in extremities None of the edge. hole of entrance c. smooth walled tunnel. stigma, lying immediately long, hole; No. 36, within a tuber single cavity b. entrance girdle chamber; the lower along of a diagram apex; VOL. as ses- found 80-130 Hydnophytum, (pollen |xm across). Beccari's new The present and flower original and species species two here reduced are that all reported had species author collected all the species a to one. in 1985 and All material is cited, marked structure. brackets afterFlower in the descriptions Smith (1967) described chambered tuber inhabited by was if n.v. able not to study seen. The a ants. the tuber figure in is the number of flowers dissected. SQUAMELLARIA Squamellaria Becc., 4 (1891) 123; A.C. Smith, Malesia 2 A.C. Smith, Fl. Vit. Nova mecodia imberbis A. Epiphytes. J. 4 Arn. (1988) cylindrical, blackish, occasionally with prominent, grey to numerous, or 36 — Engler & Prantl, Nat. (1955) 289; S. Darwin, Type species: S. imberbis numerous rings to oblong. numerous Pflanzenfam. Allertonia (A. Gray) 2 IV, (1979) 35; Becc. [Myr- (1858) 42], Spines simple, slender, mostly around entrance holes. Surface smooth, white tubercles around the tuber, to 4 to mm 2 mm in diameter. Entrance in diameter, conical, or with white, spine- fringed rims. Pores and ridges absent. Stems few scattered woody, branching 245. ovoid flexible, scattered, sometimes more holes in discrete regular K. Schumann in Arbor. Proc. Am. Acad. Arts & Sci. 4 Gray, Tuber (1886) 228; or or arising not, to 50 from cm one or long, more 1.5 cm basal stock(s); subherbaceous thick, rounded, smooth, to mostly M.H.P. Jebb: Leaves lacking spines. clustered A revision mm, the splitting opposite petiole at to with 4 Bracts lobes. Corolla towards middle, markings quadrangular the corolla lobes, on wards base of tube, pate, 110-163 occluded in diameter, pm pm; vesicles absent. Ecology species a ridge High - on 2a. at grain, Petiole 0.4-3.5 Iridomyrmex TO THE SPECIES to margin irregularly undulate. Stems 0.5 cm, Tuber apex blunt. Stems depressed at Tuber apex Leaves apices. 6-30 apex, tetra- dense forest from 200-2000 to OF Stems to rhomboid, thin, with laminaobovate flat a 1. S. or ovate markedly to margin. imberbis cordate, thick and fleshy, thickened towards their apices in middle, with 1.5 to cm in diameter cordate, apex ovate to acute. 0.8 to cm 2 their near acuminate. minutely red and green in clusters, arising m. SQUAMELLARIA arising individually, parallel-sided tapered. to- less hooked, more or markings 2. S. b. or red partially coarse, siccate, markedly or Leaves obovate, the apex rounded and sometimes Corolla lobes open, thickened towards their apex Petiole very short, apices. or green ants. lamina lanceolate cm, not with or base at sometimes reticulation fleshy entire mm, plates (squamules) indistinct lobes, Fruit margin. In- petioles. the abaxial surface. inhabited by are Stems slender, b. fringed ± with 4 the as 3.5 to exserted. Pollen 3-5-col- mm, pm; low in host tree, in open or KEY la. 4 barbate long as cordate, cm, sharply or ovoid, orange-red; calyx remains prominent. Pyrenes 4, or sometimes with All almost Stigma capitate, recurved and sometimes gonal colpi wall extensions; walls 9-14 by pollen 2-6 distorted, curving pubescent without; 6.5 first, becoming pedunculate at inflated, white throughout, or x thin, papery, homostylous. Calyx commonly lumen. Anthers 4-8 occluding 17 - between the tearing soon in axil, sessile Flowers inconspicuous. triangular 1.5 x thick, fleshy. Stipules first, florescence solitary, laterally displaced with age. Lamina 3.5 or remote. rhomboid-lanceolate or obovate, thin 55 of Squamellaria in diameter Corolla lobes obovate, major near pale their green 3. S. thekii 1. imberbis (A. Squamellaria imberbis Squamellaria Am. Arbor. Smith, Fl. & Sci. 4 Hook. Benth. f. Vit. in & Squamellaria Benth. 62266 wilsonii J. Am. nud.; 200. — 247. ex — Seemann, f., Hook, Malesia Vit. PI. 2 111. Fl. Ins. Drake, 2. Fig. (1886) 228, Fiji Myrmecodia Fl. Gen. 2 PI. Parham, — (1866) Isl. imberbis 138. — Pac. 46, figs. 1-12; ed. 2 A. 132. (1890) — 200. A.C. Am. Acad. inermis A. Hydnophytum — Smith, J. (1972) 293; A.C. Proc. Gray, Myrmecodia sphalm. (1873) Mar. t. (1964) 209, Type: Fiji, Arts Gray ex imberbe U.S. Expl. (holotype US; isotype K). (Home Arbor. Home J.W. (1988) B.C. & Hook.f. (1988) 247, figs. (1890) 4 Nov. sphalm. wilsoni; nom. (A. Gray) Becc., (1955) 289; (1858) 42; Exped. U.S. Smith, 36 Becc. Gray) S. 36 Darwin, 9A-D & ex J.W. Allertonia 13a. Baker, Type: Fiji, Baker) Becc.,Malesia ex (1955) 289; — Soc. 2 (1886) 229, PI. Fiji (1979) 35, fig. Hydnophytum J. Linn. Home 2 Parham, Bot. wilsonii 20 1139 (holotype K). Isl. 3D; A.C. Home, (1883) 365; t. 46, figs. (1964) 209, ed. Smith, A Year Drake, in 13-21; 2 Fl. A.C. (1972) 293, Vit. Nov. 4 Fiji (1881) 263, 111. Fl. Ins. Mar. Pac. 56 BLUMEA VOL. 36, No. 1, 1991 Wise. Rosemary by Drawn 467). Jebb (all pyrene d. fruit; c. flower, b. plant; Whole a. Becc. Gray) (A. imberbis Squamelari 2. Fig. M. H. P. Jebb: Tuber in liptic glossy longitudinal section, conical, 0.6 to green 4 tapering veins c. to 5-7. apiculate, a to pedunculate, branching, 1.5 x 8 5 - 3.5 to long cm and 0.3 mm, mm, pubescent without, especially Corolla tube mm 2 a long, finely more or acute, pm the foot across, diately 3 to pm, layer the Ecology given Notes unarmed late this in fruit, and - The holes is and from the wilsonii to was the tube, limb 470 vagaries of Smith to Navitho between J.W.P, P.B.T. 8326 1787 Lambasa, to pinkish; long, mm sessile pm 32 to rarely glabrous. so, 6.5 triangular, in diameter; of the to [11], Calyx quadrangular, base. Lobes grain; to with colpi walls 9-10 pm these lumina 6-16 perforations. Stigma imme- of short hairs below. Fruit fringe mm 3.5 long, blue-black; filaments height including calyx. Pyrenes several solitary, 5 mm specimens usually present 16935 longer more acute 16° 481 50' — Vanua gradually light more attenuate globose, (Beccari, of further collections and proved insignificant, SE 16° Jebb 467 471 (BISH, K), 180° 179° 468 178° or due to py- 1886), study of developmental Wairiki, 00', on Smith 564 (n.v.). Smith 750 of Mt Manuka, road to 19', Thakaundrove, 25' Korotini (LAE, SUVA), (BISH, FHO, K, SUVA), Ndelaikoro, and 30' 18° 04 472 — Devoeux Peak, Jebb Freehold uncertain. DA 11901 Seemann s.n. (n.v.). Forest (K, P). Mathuata River road near new 179° 58 Range, Western Qacavulo Estate, A.C.S, Smith 8191 479 (GH, L, UC), (BISH, K, SUVA), Above Creek, Yanawai (BULOLO, K, SUVA), 16°48 Above Wairiki, Range, Crest of Korotini (SUVA). (BO, H, UC). east of 469 Taveuni. (BISH, K, SUVA), 482 (LAE, SUVA, UPNG). Locality its shorter corolla, lobes than filaments, fruit and In the Levu. (BRI). Slopes (n.v.). Nggarawalu, (type). tuber, with fine-rimmed The thin, lanceolate leaf and flowers also iso- (BO, GH, K, UC). and Mt Somosomo (K, L, UC). wilsonii). 62266 Pass D.K. (BISH, SUVA), 16935 7 to drying. (A, BRI, CANB, SUVA), slopes to green to Flowers densely formerly distinguished by stamens (Smith, 1988). FIJI ISLANDS. - Kasi, Savusavu between cm. slightly compressed diagnostic. hooked, and leaves Collections Mt 0.1 on curved. Not m. regular, oblong, longer calyx region, to mm a most- elsewhere rare Inflorescence Corolla white, the high, cm rhomboid, apex acute, layer regularly reticulate, fleshy, fresh material, these differences have or caducous. 110-152 (134.5) as 0.4 from the remainder of the genus. species relative stages, cm, base, tapering outer not tuber, to pink. Stipules thick. Bracts strongly Altitude 300-700 - entrance less renes 0.15 to upcurving; tree. Squamellaria longer to green long to prominent below, pale corolla lobes, bud at to towards the apex. Internodes up within the lumina with minute turbinate, strongly quadrangular, a pendent lanceolate cm, Entrance holes protrusions. close very Anthers 4-5 as cm, remote pubescent. on in bud. 0.6 x wide; surface cm surface; rims upper above anthers, with 4 indistinct lobes and long, tighdy appressed in cm less recurved thickened border, about slightly to (3-)4(-5)-colpate, thick, the inner wall 3.5 pale mucronate, tetragonal wide. Pollen mm x cm, 2-3 or 50 to small hori- nearly from the side, el- and 12 high, cm on more numerous on small central process 0.6 entire mounted mid green. Midrib petiole, 18 long, cm cm Spines only present Petiole 0.4 with 30 2 subovoid to bluntly hemispherical of the tuber, less apical part when sterile. Leaves. Lamina 4.5 stem. base the on long cm Spines in diameter, cm to 57 of Squamellaria regularly oblong less more or to grey. Stems several, ly arising to black. to matt revision from host, growing horizontally zontal below, swollen above, A Horne Prov., 480 Nggathavula Estate, 1139 U.S. (K) (type DA of S. Expl. Exped. US 58 BLUMEA VOL. 36, No. 1, 1991 Wise. Rosemary by Drawn 483). Jeb b—d: 475; Jebb (a: fruit d. bud; c. flower, b. plant; Whole a. Smith, A.C. major Squamelari 3. Fig. M.H.P. Jebb: A 2. Squamellaria major Squamellaria major ed. A.C. 2191043 Smith, A.C. (1972) 293; 2 A.C. Smith Fl. Vit. Nov. cm cylindrical, obovate, in diameterwith Stems several, nodes 0.5-1.5 cm, Lamina obovate near from to 20 narrowed 7 to inflated towards middle, to 2.5 at to 0.5 at the pollen grain wall, to colpi being with 4 or 3 x - pale a red pm, the foot 17 x to pm 6.5 0.3 or on trunks The leaves of this Collections - 8323 483 5 to at mm Tuber fusiforme, incrassati orientes. the centre, paralled when Folia 16° tumidum apicem to mm. Corol31 about by edges 6 pm, the a pm in diameter; long as the as number of fine of the outer mm Tube mature. towards apex, and then 125-163 (140) across, or wall mm; colpi height of outgrowths of and colpus interdigitate; irregularly reticulate, small numerous large limbs of trees in closed forest species always 180° out- to 3 mm. to Pyrenes hooked, at 300-400 m. appear chlorotic and diseased. Exami- 00', cover the out- in them. SE on 16° 50' road to Jebb Jebb, spec. inter versus, fasciculata, cotypus 50' thekii ovoideus, calyce BISH, K; 35 above the disc. Anthers 5-6 prominent calyx fungal hyphae margine corrugata. Calyx Fructus to long. 179° 59' Slopes Devoeux Peak, of Mt Jebb Manuka, 475 east of Wari- (FHO, SUVA), 476 (SUVA). Squamellaria paullo apices. at Margin irregularly with FIJI ISLANDS. Taveuni. (type). (BISH, SUVA), 3. entrance few in number. side of the tuber have Smith cm across. apex rounded, cm; nation of fresh tuber material reveals that the small white tubercles that ki, Entrance of ring sessile. Flowers [3] within the lumina with layer Surface cm. long. cm each venation branch, often at narrowing less occluded to x pink patch or mm 5-colpate, 14 17 - gentle undulations, which arise from both Growing - solitary Note US = mm. Ecology Often Isl. Fiji smooth, grey; inter- above and below. in diameter, to more or fleshy, quadrangular below, 5.5 PI. 8323 square in section with 4 mounds, exserted above anthers. Fruit growths. Stigma to surface rounded in bud, sides 12 pm thick, the inner wall these lumina 9-30 cm, Smith cordate, drying thin, yellowish or Inflorescence cm. mm Pollen 4- mm. grain, 1 7 to 4.5 x blunt mouth; barbate scales 2 the walls 8 prominent margin, barely thickened, colpi the 1.5 to within each fleshy, cm, narrowing abruptly pubescent, mm, widening slightly 1.5 x la recurved, white, lobes green with lobes 24(-40) to flexible, to immediately abruptly narrowed, Lateral veins apex. Calyx cup-like, apex blunt broadly oblong-elliptic, to undulate, white. Petiole 0.3 edges Parham, Type: Fiji, — shortest and thickest towards apex. Leaves clustered not, base or filaments J.W. (1967) 90; 247. thickened, spine-topped, white rim a green above and below. Midrib long; 37 (1988) stiff Spines apparently arising holes, unbranched, scattered, bifurcate 4 (holotype US; isotype BISH, K, L, US). Tuber subpendent, apiculate 3. Fig. — 59 of Squamellaria Cont. U.S. Natl. Herb. Smith, with small white tubercles, black. holes 1 revision annulos ad obovata 8 mm usque manifeste — Fig. foraminum, ovata, coronatus. apice alba, — lobi 4. qua foramina diametro, ramosi, 4-dentatus. Corolla 478, SUVA). nov. acuto ab tubere usque tumidi in Typus: Fiji, margine uno rotundo, alabastro, Jebb 477 incrassato. fasciculo basi Caules vel pluribus obtuso, carnosa, obovati in maturitate. (holotypus SUVA; isotypi 60 BLUMEA VOL. 36, No. 1, 1991 Wise. Rosemary by Drawn 477). Jebb (all pyrene f. apex; fruit at disc and calyx e. fruit; d. bud; c. flower; b. plant; Whole a. thekiJeb . Squamelari 4. Fig. M.H.P. Jebb: Tuber surface 5 cm 1.5 numerous Stems several, across. 36 0.8 x a Stipules rounded, Bracts papery, to 0.2 less to cm. 0.2 to one to long, 9-15 rise giving wide; cm hair-like to in numerous, the inner edge cm, few base and apex, ab- to cm some than 0.1 to lobes, 31 to Corolla tube mm. ments 8 to mm, unbranched, to 3 Pollen 4 mm. ened; walls to 14 pm, these lumina9-22 anthers, 2 to mm colpate, colpi 4.5 pm, the to round the foot deep layer in diameter, on groove the above the disc. Anthers 150-163 pm; the inner wall pm, a mm 19 pm to wall outer 3.5 across, to 6.5 x 3 fila- mm, thick- edges irregularly reticulate, within the lumina smooth. slightly recurved, mm, on gently surface, outer to with 10 to mm, abaxial prominent a above Stigma in diameter, rounded, margin reflexed and fimbriate. Fruit ovoid, fleshy. Pyrenes to 0.2 fleshy, quadrangu- minutely pubescent 6 to to sessile. Inflorescence mm, Corolla white, with mm 5 to Calyx towards middle, less orbicular in bud, scales 2 more or mm. margin. at swelling recurved. Lobes obovate, blunt, 36 to 0.8 apex. Lamina cordate at caducous. long, cm to 1 rings abrupt, scattered clusters, Leaves clustered cm. Flowers [2] 4-dentate, abruptly flattened lar, protuberances, from arising 27 to black. Entrance holes to thickened rim Internodes 0.3-3 cm. holes, rhomboid, apex acute, base blunt to cordate, margin crinkled. Petiole up ovate cm. 61 of Squamellaria entrance small dark brown long, cm of rings with apart; prominent, cm to with rough, to spines revision subpendent, regular, cylindrical-fusiform, tapering swollen between the ruptly A keel. Ecology 400 Notes gest an low level epiphyte, growing species status. in S. species; clusters The two species, only trunks of on at stems and S. imberbis the Its stem trees in forest at 300- apparently the tuber surface is stems are not as com- and leaf characters sug- but flower characters indicate species, on and once, sympatric. between the other The distribution of major collected with which it is intermediateposition specific peculiar to this scattered, while here they arise in the apex of the tuber and elsewhere. specific epithet derives from the Fijian name 'theke theke nkau' for ant-plants, 'testicles of the trees'. meaning Collections. 477 A A little known - the mon as its - m. (type), 478 FIJI ISLANDS. Taveuni. SE 16° 50' 180° 00', Taveuni road to Devoeux Peak, Jebb (cotype). ACKNOWLEDGEMENTS I tion, thank F. White and C.R. and Rosemary Huxley Wise for her for their support, meticulous A.C. Smith for nomenclatural clarifica- drawings. REFERENCES BECCARI, O. 1886. Piante HUXLEY, C.R., the JEBB, Hydnophytinae. M.H.P. Thesis, SMITH, SMITH, 1985. Oxford A.C. from Fiji. A.C. Ospitatrici. Malesia 2 : 228-230. & M.H.P. JEBB. 1991. The tuberous Blumea 36: Taxonomy 1-20 and epiphytes of the Rubiaceae 1: A new subtribe - (this issue). tuber morphology of the rubiaceous ant-plants. D.Phil. University. 1967. Studies Contr. U.S. 1988. Flora of Pacific Natl. Herb. Island 37: plants XVIII. New and noteworthy flowering plants 69-106. Vitiensis Nova 4. Pacific Tropical Botanic Garden, Hawaii.