A. Hay, 1988 25

Anaphyllopsis: A New Neotropical of -Lasieae

Alistair Hay Department of Sciences South Parks Road Oxford England

During the course of revising the spe­ phyllous (under normal conditions of cies of in the Far East, it growth) in C. americanum, whereas in became necessary to review generic limits it is rhizomatous and pol­ in the tribe Lasieae sensu Engler ( 1911 ) as phyllous, and in it is cormous amended by Bogner ( 1973 ). It has become and monophyllous. apparent that the pantropical Cyrtosperma It is proposed here that a new genus, circumscribed by Engler (loc. cit.) is , be erected as an alternative heterogeneous. to "lumping" Dracontium, Dracontioides Three species of Cyrtosperma have and C. americanum. Were the latter been recognized for the New World. Two course to be adopted, the resulting broad of these belong in extant genera--C. generic concept of Dracontium would be wurdackii Bunting (Urospatba) and C. inconsistent with the existing rather nar­ spruceanum (Schott) Engler (Dra­ row limits between other genera of the contium). The necessary new combina­ Lasieae such as , Urospatba, tions are to be made elsewhere, in a and Cyrtosperma s.s. forthcoming revision of Cyrtosperma. The Two new species of Anapbyllopsis are third species, C. americanum Engler, can­ described, both, sadly, from single frag­ not be fitted into any presently recognised mentary collections. Their leaf blades genus. however, are so distinctive as to justify drawing attention to these as repre­ Subtribal and generic limits in the sentatives of new and apparently rare Lasieae are also to be dealt with elsewhere. species. However, the species in question differs from Cyrtosperma s.S. in the presence of both prophylls and cataphylls on the Anapbyllopsis A. Hay, gen. nov. rhizome, and in having fenestrate foliage. Herbae terrestres paludosae; caudex The absence of cataphylls may be used as a hypogaeus rhizomatosus, prophyllis diagnostic character for the subtribe La­ cataphyllis euphyllisque praeditus, siinae which includes the. Far Eastern orthotropus; folii lamina fenestrata species of Cyrtosperma and in which vel pinnata; spatha spiraliter torta; fenestrate leaves are unknown. Fenestrate flores hermaphroditi tepalati; sem­ leaves are present however, in Dracontioi­ des and Dracontium. Cyrtosperma ina campylotropa albuminosa; testa americanum appears to be more closely crassa ornata. related to Dracontioides and Dracontium, Solitary herbs with subterranean but differs from both in the form of the orthotropic rhizomes (where spathe which is spirally twisted rather like known) bearing prophylls, cata­ that of the Indian Anapbyllum wigbtii phylls and euphylls; leaves generally Schott in C. americanum, and hood­ solitary with unarmed tubercular to forming to erect in the other genera. Of the smooth often mottled petioles; lami­ vegetative characteristics, the stem is nas dissected by fenestration or rhizomatous and the sympodium mono- ternate with fenestrate segments or 26 AROIDEANA, Vol. 11, No. 1

simply pinnate; termi­ Stem unknown; leaf solitary; petiole nal, solitary, on long peduncles simi­ mottled, warty, ca. 1.5 m long, ca .1 lar to the petioles; spathe mem­ cm diam. (dry); lamina pinnate; branaceous, papery when dry, lowermost leaflets (= posterior marcescent, convolute towards the lobes) stalked for ca. 3 cm, their base, spirally twisted in the mid and blades ca. 30 cm long, 11cm wide, upper part; spadix with a stipe oblong-ovate, more or less symmet­ adnate for most of its length to the rical, with the tips acuminate for ca. spathe; flowers hermaphrodite 2 cm; primary lateral veins (of throughout the length of the spadix, leaflets) pinnate, ca. 12 on each side tepalate, maturing (like the fruits) in of the midrib, brochidodromous, a basipetal sequence; perianth and collected into a more or less undu­ androecium tetramerous; stamens lating sub-marginal vein, not promi­ with short, strap-like filaments and nent below (in dry state ); secondary extrorse bilocular anthers opening venation reticulate, faint; anterior by apical pore-like slits; pollen rachis ca. 13 cm long, with two pairs ovoid, monosukate, with more or of opposite leaflets similar to basal less smooth exine; ovary superior, pair but progressively smaller, ter­ unilocular, uni-or biovulate with minating in an incompletely divided basal placentation; stigma not nec­ pair of segments; peduncle smooth tariferous; fruit ovoid to obpyrami­ otherwise similar to, and winding dal, indehiscent; seed campylotro­ round the petiole; spathe membra­ pous, pachychalazal, albuminous; nous and papery (dry), ca. 16 cm seed-coat thick, verruculose to long, ca. 7 cm wide at widest, deeply channelled. Type species: A. ovate-Ian ceolate (flattened), convo­ americana (Engler) A. Hay: (ba­ lute in the lower quarter, spirally sionym Cyrtosperma americanum twisted in the upper 2/3 greenish Engler). Distribution: Tropical South outside, reddish brown within; spa­ America - , Surinam, dix 2.3 cm long, 8 mm diam., and . Widely stipitate for ca. 6 mm with the stipe dispersed, the collecting sites of the adnate to the spathe; flowers tetra­ three species (Fig. 4.) fall within merous; ovary unilocular; ovules three of the Pleistocene forest refu­ two; stigma raised for ca. 1 mm on a gia proposed by Prance ( 1982). conical style; fruit and seed un­ known, Fig. 1. Type: Venezuela, Territorio Amazonas, Rio Negro, Key to species of Anapbyllopsis near the Brazilian border around 1. Lamina pinnater; leaflets ovate, petio­ Piedra de Cucuy (approx. 66 51" W; lulate ...... A. pinnata 1 13" N), 2 Nov. 1968,]. G. Wessels 1. Anterior lobe pinnatifid, or if inter­ Boer 2387 (U! holotype). Distribu­ preted as pinnate then the leaflets sessile tion: known only from the type with broad insertions ...... 2 collection from tropical rain forest 2. Lateral segments of anterior lobe with on granitic soil; altitude 100-350 m. retuse to bifurcate apices ...... Anaphyllopsis americana (Engler) ...... A. americana A. Hay, comb. nov. 2. Lateral segments of anterior lobe with Cyrtosperma americanum Engler in rounded apices. . . . . A. cururuana Martius, Fl. Bras. 3,2 (8178) 117, t. 22 & in DC, Monog. Phanerogam. 2 (1879) 272 & Anapbyllopsis pinnata A. Hay, sp. in Pflanzenr. 48 (IV. 23C)(1911) 22, fig. 8; nov. Pulle Enum. Vasco PI. Suriname ( 1906) 78; A ceteribus speciebus Anaphyllopsidis Jonker-Verhoef & Jonker in Acta Bot. folio veriter pinnato differt. Neerl. 15 (1966) 130. Type: French Gui- A. Hay, 1988 27

'A Fig. l. A. americana mottled green, grey and brown, weakly pustular, unarmed, spongy, ana, Leprieur 152 (P! lectotype, selected conspicuously geniculate at apex; here; US!) sheath very short, rarely to 8 cm, Herb to 1.5 m tall; rhizome straight, membranous; lamina dark green, subterranean, orthotropic, brittle, glossy above, dull below, sagittate; ca. 2-3 cm diam., to ca. 15 cm long anterior lobe pinnatisect by fenes­ with few scattered roots (in anero­ tration (juvenile expanded leaves bic conditions); or short with copi­ sometimes remaining fenestrate ), ous roots (aerobic conditions); leaf lateral segments opposite, separated solitary, rarely to three together; to near midrib, unicostate, with petiole to l.5 m long (usually less), submarginal veins, costae divided 28 AROIDEANA, Vol. 11, No.1

and running to the extremities of the ca. 6cm wide at widest point, retuse or bifurcate tips; terminal lanceolate, convolute below, segment rhomboid; posterior lobes twisted above, erect, pale greenish­ naked in the sinus for ca. 5 cm, brown to reddish outside, ivory with entire on the posterior side, more or purpose margins below within; spa­ less pedately divided by fenestration dix on a stipe completely adnate to on the anterior side; inflorescence the spathe and thus displaced at an solitary; rarely with an inflorescence angle to the peduncle, ca. 2.5 cm and infructescence present together long, ca. 7 mm wide; flowers tetra­ on one plant; peduncle slightly merous, ovary unilocular; stigma shorter than and similar to the dry, button-like, very small; ovule petiole, present with leaf; spathe solitary, with basal placentation; (and spadix) drying black and turn­ fruit obpyramidal, ca. 1 X 1 cm, ing black in alcohol, to 18. cm long, flat-topped and closely spaced form- Fig. 2. A. pinnata A. Hay, 1988 29

ing channels in between, Fig. 2. the structure of the seed. Engler's illustra­ Distribution: Surinam and French tion (loc. cit.) gives the impression of a Guiana; growing in lowland swamp thin testa surrounding a thick runinate forest, swamp forest regrowth, endosperm enclosing the embryo. This is swampy woodland and scrub. not so. His "embrog" is endosperm con­ I observed this species in August 1984 taining the embryo. His "endosperm" is growing in and near the ORSTROM forest the seed coat, and his "testa" is the regeneration plots along the Piste Ste. Elie seed-coat's outer pigmented layer. The near Sinnamary in French Guiana. Plants channelled petiole and the inflorescence were growing in the shade of swamp forest emerging without a leaf from the mounts floor, and were rare, in individuals widely of two cataphylls are at best atypical, at scattered. They were found to have the worst fictitious. rhizome below the water table, with the Specimens seen: apical bud situated at the interface of French Guiana, Saut Mais ± 20km E. of slate-grey waterlogged (ferrous) and the Saul, Cremers 6107 (CAY!, K!); overlying brown (ferric) aerobic soil. The Piste Ste elie, nr. Sinnamary, Hay bases of the rhizomes lay level with the top 2818 (M!); Leprieur 152 (P!, US!), of an underlying layer of sand. In such s.n. leg. 1838 (P!), leg. 1847 (P!); conditions the rhizomes bore very few Marecages de Mt. Sineri, Melinon roots, scattered along the length. Some of 52 (P!), 98 (P!), 115 (P!). these were wrinkled, suggesting they are Surinam, confluence of the Ta­ contractile. A few plants were also found panahoney and Paloemeu rivers, growing in an area flooded by a stream blocked off by the road which had been Wessels Boer 1223 (Kl, NY!, U!, built eleven years previously. A consid­ US!). erable depth of sand had been deposited Anaphyllopsis cururuana A. Hay, by the stream over the forest soil, the trees sp. nov.. had died and the canopy had opened up. Ab alteris speceibus Anaphyllopsidis Here A. americana was much more vigor­ lamina ternato-pinnata, lobis posticis utrin­ ous than when growing within the forest, que fenestratis vel sectis, fructo ovoideo, some with three leaves at once and two differt. . The water table was more Stem unknown, ca. 30 cm below soil or less level with the sand, and the canopy level; leaf solitary; petiole ca. 1.3 m had opened up. Here A. americana was long" mottled green and white, much more vigorous than when growing weakly tubercular, unarmed; blade within the forest, some with three leaves at tripartite; posterior and anterior once and two inflorescences. The water lobes stalked; anterior lobe pinnati­ table was more or less level with the sand, fid by fenestration, divided to or and the rhizomes had apparently grown up very near to the midrib, with the through it. They had become short, with a proximal lateral segments opposite great many roots clustered around the or subopposite, bicostate; costae crown. Curiously, although several plants were in fruit, no seedlings or juvenile divided shortly above junction with plants were found at this site, whereas they midrib; apices of the lateral seg­ are present in the forest. In areas of free ments narrowing to rounded, standing water, a few plants were found slightly asymmetrical and oblique growing at surface level in the corner of apices; distal segment broad, if di­ buttresses of dead boles, rooted in moss, vided then more shallowly than the bark and mud collected there. Part of the proximal segments; posterior lobes material I collected was lost in transit to pinnatifid by fenestration or the United States, and my collections are fenestrate on both sides of the represented by some living plants kindly posterior midribs; tips of the poste­ received at Munich by ]osefBogner. I have rior lobes acuminate for ca. 2 cm; therefore been unable to examine in detail inflorescence solitary on a peduncle 30 AROIDEANA, Vol. 11, No. 1

Fig. 3. A. cururuana Alto Tapajos, Rio Cururu, 7 Feb. 1974. W. R. Anderson 10627 (US! similar to the petiole; spathe mem­ Holotype; NY!, lAP Isotypes). Distri­ branous, spirally twisted in upper bution: known only from the type part, convolute in the lower, to 18 collection, in water at the edge of a cm long, marcescent; spadix ca. 5 canal at ca. 140 malt. cm. long, 1.2 diarn. (intl.), ca. 8.5 cm long, 2.5 cm diarn. (infr.), stipitate for ca. 8 mm; stipe completely Judging from the herbarium sheets, the leaf of this species is first branched (at the adnate to the spathe; flowers tetra­ top of the petiole) in the strict sense of merous, ovary unilocular, uniovu­ branching by outgrowth, and thence fur­ late with basal placentation; fruit ther divided by dissection within the ovoid, smooth; seed orange-brown; margin of the lamina (fenestration). It is testa irregularly crested, brain-like, thus intermediate in this respect between Fig. 3. Type: Braxil, Estado de Para, A. americana, in which dissection of the A. Hay, 1988 31

never fenestrate, suggesting marginal out­ growth of the lobes of the blade. It would also appear that in some Dracontium species there is both marginal and in­ tramarginal dissection in the same leaf.

ACKNOWLEDGMENTS I am indebted to Marie-Francoise Pre­ vost and Hubert de Foresta for kind hospitality and assistance in the field in French Guiana in 1984, and to Josef I \ Bogner for skillfully growing Anaphyl­ . ' lopsis americana at Munich. I would also like to thank the curators and directors of the following herbaria for allowing me to examine material held in their institutions: CAY, K, NY, P, U and US. I / I This work formed part of a thesis ~ Fig. 4. Distribution of I' collections of Anaphyl­ submitted for the degree of Doctor of lopsis species, mapped , Philosophy in the University of Oxford. I over Prance's (1982) i am grateful to my supervisor Dr. D. ). __ ~~ distribution of proposed Mabbecley for his guidance and support. I ' leisticene forest refugia ' I am grateful to Rosemary Wise for I in South America. / illustrating the species.

blade occurs within the margin, and A. REFERENCES pinnata, in which the leaflets have the Bogner,). 1973. Die Gattung appearance of having being formed en­ Pycnospa­ tirely as outgrowths of the leaf primor­ tha Thorel ex Gagnep. (Araceae). dium. The leaves of this genus seem very Osterr. Bot. Z. 122: 199-216. interesting developmentally, and are in Engler, A. 1911. Araceae-. need of investigation. The occurrence Pflanzenreich Heft 48 (IV. 23B). together of two such seemingly opposed Engelmann, Leipzig. methods of leaf dissection within one Prance, G. T. 1982. A review of the genus may be matched by Amydrium phyteographic evidences for Pleisto­ Schott (Monsteroideae). Juvenile leaves of cene climatic changes in the Am. medium are fenestrate, while those of Neotropics. Ann. Mo. Bot. Gard. 69: Am. magnificum, though dissected, are 594-624. D