36 AROIDEANA, Vol. 25 Ulearum donburnsii, a New Species of from Ecuador

Thomas B. Croat Missouri Botanical Garden P.O. Box 299 st. Louis, MO 63166-0299, USA Betsy Feuerstein Memphis, TN

ABSTRACT the Araceae. The tribe comprises the gen­ era, Zomicarpa, Zomicarpella, Ulearnm, Ulearnm donburnsii, a new species of and Pilarnm (Bogner, 1997; Boyce, 1995; Araceae, is described from the Amazon Mayo et al., 1997). All of these genera are lowlands of Ecuador along the RIO Morona relatively restricted in geographic terms. in the Province of Morona-Santiago. The Most of the genera, with the exception of species is only the second species in the Zomicarpa, are restricted to the upper Am­ Ulearnm. The species differs from azon basin along the margin of the Andes. the only other species, Ulearnm sagitta­ Zomicarpa is found at about the same lat­ tum Engl., by having filiform staminodes itudes but it is restricted to eastern . on the spadix on either side of the stami­ The genus Ulearnm is also found in the up­ nate portion of the spadix, in contrast to per Amazon basin, but this is the first spe­ the depressed globose or short, cylindric cies of the genus in Ecuador. staminodes present in Ulearnm sagittatum. Ulearnm donburnsii has blades similar to Ulearnm sagittatum but differs in hav­ INTRODUCTION ing a much longer and thinner appendix A recent discovery of a in the Am­ and in having long filiform starninodes azon lowlands in the Province of Morona­ rather than depressed, globose, or short Santiago has proven to be a new species. cylindric staminodes. Ulearnm donburnsii The new species is known from a single with its attenuated starninodia somewhat locality and was introduced into horticul­ resembles the attenuated filaments of Pi­ ture by a nursery in southern Ecuador. The larnm mansericbense Nicolson. Both spe­ species was first brought to the attention cies also share similar leaves and short­ of the senior author by naturalist, Betsy creeping rhizomes. The resemblance is Feuerstein, who subsequently revisited the however superficial since in the case of Fi­ site to acquire much needed habit and larnm most of the elongate structures are growth information. It is unusual to dis­ actually attenuated filament-like connec­ cover new species in the Amazon low­ tives of the fertile stamens whereas in lands, though the Rio Morona lies near the Ulearnm the elongated structures are fili­ Cordillera del Cutucu, an isolated range of form staminodia not directly associated low mountains at the edge of the Amazon with fertile stamens and instead arise di­ basin which has many endemic species. rectly from the axis of the spadix. In Fi­ Most of the new species in the region of larnm the prolonged filament-like connec­ Cordillera del Cutucu occur further up on tives are present on a broad and continu­ the slopes of the massif so it is unusual to ous band above the pistillate portion of find an endemic in this region at only the spadix whereas in Ulearnm donburn­ about 243 m elevation. sii the filiform staminodes occur mostly in The new species is a member of the tribe two narrow bands on either side of the Zornicarpeae in the subfamily in staminate portion of the spadix. T.B. CROAT, B. FEUERSTEIN, 2002 37

KEYTOSPECmSOF~UM

1. Spadix with depressed globose or short, cylindric staminodes below and above the male flow- ers; appendix short and thick, less than 5 times longer than wide ...... Ulearurn sagittaturn Engler. 1. Spadix with long filiform staminodes below and above the male flowers; appendix elongate and slender, more than 15 times longer than wide. " Ulearurn donburnsii Croat & Feuerstein

Ulearum do"burnsii Croat & Feuerstein, sheath, but instead the petiole bases are sp. nov. surrounded by a cataphyll 4-6 cm long, the cataphyll apex acute to rounded and Rhizoma gracilis, nivius, 6-9 mm diam.; weakly apiculate; blades sagittate-hastate, petiolus (9)14.5-23.5(31) cm longus, 3-4 8.0-14 cm long, 6.0-13 cm wide, 7.5-14.2 mm diam., teres, infirme multi-costatus; cm long from tip of blade to tip of poste­ lamina sagittato-hastata, 8.0-14 cm longa, rior lobe, matte-subvelvety, variable in col­ 6-13 cm lata; lobulas posteriorus, aequans or, ranging from nearly solid dark green or vellongior quam lobula anterior; sinus an­ mottled with light green or white above guste V-formatus vel anguste spathulatus; (those with the most solid dark green venas basales 4-6 utroque; nervis primariis blades usually with a continuous, solid lateralibus 1-2 utroque; pedunculus 14.2- pale green V-shaped band paralleling the 27(42) cm longus, 3-4 mm diam.; spatha margins), moderately paler and matte be­ anguste ovata, apertus, viridis; spadix 6- low; margins straight to undulate and of­ 7.2 cm longus, palide viridis; spadix pisti­ ten crisped and hyaline; anterior lobes lata 1.2-1.3 cm longa, 2-3 mm diam.; pis­ tilas 20-27; appendix 3.0-3.7 cm longus, 5.0-7.2 cm long, decurrent onto petiole at palide viride; stamens cum filamenta ca. 1 base; posterior lobes 5.5-9.7 cm long, 3.1- 3.5 cm wide, equaling or up to 1.2 times mm longa; bacca 6.7-7.5 mm longa, 3.0- longer than the anterior lobe, and directed 3.3 mm diam., elliptica; semina 4.9-5.3 mm long, 2.3-2.5 mm diam. Typus: EC­ toward the base or directed somewhat UADOR. Morona-Santiago: Along ruo Mo­ outward at 40-50° angle, bluntly pointed rona, along road E. of Santiago and E. of at tip; basal veins 4-6 per side, 1st pair free Rio Santiago, Amazon lowlands, flood to the base, 2nd pair coalesced 4-8 mm, plain of ruo Morona, vicinity of Puerto de somewhat retrorse then prominently Morona on E. side of River, 200 ft E. of arched toward the apex, 3rd and 4th coa­ ferry which crosses to San Jose Morona, lesced 1.5-2.2 cm, the 3rd pair usually di­ along margin of flood plain forest and rected straight to apex; sinus deeper than adjacent to the road, 02°55.341 'S, broad to about as broad as deep, narrowly 77°43. 272'W, 183 m, Croat 84834 (holo­ V-shaped to narrowly spathulate, usually typus MO-5564275; isotypus AAU, B, CAS, 0.5-2 cm wide in life, 1-3 cm on flattening, COL, F, GB, GH, K, L, M, NY, Q, QCNE, sometimes closed with the lobes overlap­ QCA, S, U, UB, US). ping; major veins etched to etched-sunken Terrestrial rhizomatous herb; rhizomes and concolorous above, darker, moderate­ slender whitish, 6-9 mm diam., sometimes ly raised and obtusely angular below; mid­ branched; petioles (9)14-23(31) cm long, rib somewhat thicker than broad toward 3-4 mm diam, terete to subterete, multi­ base, faintly and sparsely pale-lineate; pri­ ribbed circumferentially, usually obtusely mary lateral veins 1-2 pair, arising at 45- sulcate above the middle or at least toward 55° angle; higher order veins (2nd and the apex, sometimes narrowly and deeply 3rd) clearly distinct on lower surface. In­ sulcate near apex (sometimes with the florescence solitary, erect; peduncle 14.2- margins moderately acute) medium green 27(42) cm long, 3-4 mm diam., matte, with minute dark green speckles (these spongy, deeply sheathed and sulcate al­ speckles sometimes becoming purplish), most to the base, with one margin folded matte, spongy; lacking any sign of a over the other, only the apical portions 38 AROI D EANA, Vol. 25

Figs. 1--4. Ulearum donburnsii Croat & Feuerstein 1 (upper left). Habitat with . 2 (upper right). Inflorescence. 3 (lower left). Eleven pl ants from a single population demonstrating the degree o f va riati o n in leaf shape and colo rati o n. 4 (l ower right). Leaf blade showing sagittate solid green bl ade with unequal, more or less para ll el poste ri o r lobes exceeding the le ngth of the ante ri or lo be. ope n and sulcate, sparsely blotched w ith cm lo ng, 2-3 mI11 cli am. , fused obliquely darker transverse lin es; spathe narrow ly to the spathe in the lower 1;2 o r fu sed ovate , 5.8-7.0 cm lo ng, 2.0-2.7 C111 w ide, throughout (sometimes eve n with a sma ll erect-spreading, medium green to pale portio n of the steril e staminate secti o n, green o r white to green tinged pinkish along w ith the filifo rm staminodia, fu sed gra y, some times mo ttled w ith pinkish to the spathe), the lowermost pistils in a gray, acuminate at apex, decurrent at base Single row consisting of 3-5 fl owers bo rne o nto peduncle, tightly fo lded toward the in the middle of the axis (sometimes w ith base, the main veins 13, the margins in the a small gap several fl o wers above the low er 112 o f the spathe rolled under, those base), succeed ing p o rtio ns with more o f the uppe r 1;2 ro ll ed in ward, inner sur­ fl owers per row, up to 5 fl owers per row; face smooth and glossy, o uter surface with a secti o n o f filifo rm staminodia 2. 7- matte, on magnification with a minute al­ 3. 0 cm long, to ca. 1.5 cm Wide, this en­ veolate pattern o f irregular ridges and veloping the staminate secti on , occurring deep depressio ns; spadix 6-7.2 C111 lo ng, both above and below the staminate sec­ overtopping spathe by ca. 1 cm, w ith the ti on, mostly in the first 2--4 m111 above and pistillate po rtion interrupted by a segment below the staminate section but also ex­ with lo ng filiform staminodes surro und ing tending up to 6 mm above the staminate a short staminate portio n, sometimes w ith secti on and sparsely so aU the w ay to the a naked portio n of the axis just above the pistill ate section; the filiform staminodia pistilate po rtion; pistil ate portion 1.1- 1.8 ca. 20-25 in all , greenish, 0 .6)6.2-7.0 mm T.E. CROAT, E. FEUERSTEIN, 2002 39

Figs. 5- 8. Uleanun donburnsii Croat & Feuerstein 5 (upper left). Leaves with subhastate blades with a paler green band extending along the entire border. 6 (u pper right) Leaves with subhastate bicolorous blades with pale green areas between the majo r veins. 7 (lower left). Infructescence with the persistent spathe fl exed forward. 8 (lower right) . Short-creeping rhizome with sparse roots. lo ng, 1-2 mm diam., terete to somewhat some\vhat thicke r in the distal half, w ith fl atte ned , minutely puberule nt, drying basal placentation; funicles 0.6- 0.7 mm with weak ri bs, these finely and minutely long; both the funicle and ovary moder­ warty/ papillose; the axis of the staminate ately fl attened ; stamens usually with the spadix as well as the fil ame nto us p rocess­ fil ame nts ca . 1 mm long, rarely w ith the es w ith w hitish-drying, elo ngate cellular stamens borne on branched fi laments; an­ inclusions 0.1-0.4 mm lo ng; staminate spa­ thers with a pair of thecae 0.8-1.0 mm dix (including appendix) (5)6.6-7.5 cm w ide, yellowish green, truncate and irreg­ long, usually fa lling off (or eaten ofo after ula rl y ro unded at apex bu t weakly con­ anthesis; stal1l.inate fl owers arranged in Lf- nected to appear like the fi gure 8, each 5 weak spirals or rows; pistils 20-27, l.0- theca w ith a medial pore (this 0.1 mm 2.8 mm lo ng, 0.8-1.0 mm diam , cylindri cal d iam.) , drying w ith a pale r, narrow ly to somewhat swollen, somewhat oblique raised edge (also on dly ing appear to be at the base, the surface weakly sho rt pale­ discrete and separate anthers with medial striate; stigma 2 mm wide round, funnel­ pore rather than individual thecae); poll en shaped with a deep central depression, ovoid , 20-22 fL , echinate, is of medium sometimes somewhat obli que; ovary 1- size and is p resented in mo nads, the locular w ith a short style; stigma discoid spines 1.5 fL long; appe ndix 3.0-3.7 cm and sunken in the center; ovul e 1, anat­ lo ng, mo re o r less flattened, 1 X 2 mm ropous, subcylindric ovules 0.9-1.1 mm cli am. , pale green, ca. 1. 5 cm lo nger than long, 0.5-0.7 mm w ide, 0.2 111m thick, and the spathe. Berries 6. 7-7.5 mm long, 3.0- 40 AROIDEANA, Vol. 25

Figs. 9-12. U!earum donhurnsii Croat & Feuerstein 9 (upper left). Leaf blade showing pattern of venatio n and variegations in mottling with three shades of green. 10 (upper right). Base of uprooted plant showing roots, stem, petioles with mottled bands and unopened leaf contained in leaf sheaths. 11 (lower le ft). Leaf in bud enveloped by mottled leaf sheath. 12 (lower rig ht). Infru ctescence w ith re fl exed spathe and exposed berries. T.B. CROAT, B. FEUEHST EIN, 2002 4]

Figs. 13-18. Ulearum donburnsii Croat & Feuerste in 13 (upper le ft). Anthe r with central po re and eme rging po ll e n g rains. 14 (upper ri ght). Clu ste r of po ll e n grains. 15- 18. (Middle and lower right). Poll e n grains in va ri o us states of magnificatio n indicated by bar.

3.3 mm cliam. , ellipsoid to e llipsoid-ovoid , sessile and bo rne at the base of the ova ry, whitish in lower 1!J , pale yellow-green in testa thin, faintl y brownish white; raphe upper 213; pe ricarp thin, mottled irregularly inconspicuous; embryo large, pale yell ow with pale purple (mottling \N ith jagged green, straight, attached basally by a thi ck­ edges); seed solitary, 4.9-5.3 mm lo ng, ened whitish funicl e less than 1 mm long, 2.3-2.5 mm cli am. , appearing essentia ll y endosperm lacking. 42 AROIDEANA, Vol. 25

Ulearum donburnsii is known only zornes of these genera may be short­ from the type locality in tropical moist for­ creeping and tuber-like, investigations est in Morona-Santiago Province of eastern made by Wilbert Hetterscheid (pers. Ecuador at 283 m. Although there is a pos­ comm.) with Filarum are instructive. His sibility that Zomicarpella occurs in Colom­ observations demonstrated that "segments bia, this occurrence of a member of the of tlle rhizome showed that the apical bud Zomicarpeae in Ecuador is the northern­ for the new growth was clearly excentric most collection for any member of tribe on the stem, that there was a narrowed Zomicarpeae from the Andean region of part on one side of the older part of the western South America. The genus Zomi­ rhizome, and that the fresh roots were lo­ carpa is restricted to eastern Brazil while cated on only one side of the rhizome". Zomicarpella is known for certain only Accordingly all of these features prove that from western Brazil. The range of Ulear­ the stem is rhizomatous rather than tuber­ um previously known from eastern Peru ous. Even Zomicarpa, based on observa­ and western Brazil is extended to southern tions by Eduardo Gon~alves (pers. comm.) Ecuador. The species may be more wide­ may have tuber-like rhizomes like those spread than is currently assumed because described above by Hetterscheid. If so this when not in flower the are not would mean that all of Zomicarpeae have markedly conspicuous and could easily be rhizomatous stems. confused for sterile juvenile plants of some The species is named in honor of our other genus, especially Syngonium. good friend, the late Don Burns, who was According to Michael Hesse (Univ. of a grower of many Araceae, and an active Vienna) the pollen of utearum sagittatum, member and officer of the International the only other species in the genus, differs Aroid Society, whose computer skills were in being more spherical and larger (26-28 helpful to many. Don was for many years microns) with longer spines (to 2 IL). Ad­ responsible for maintaining membership ditionally the pollen of U. donburnsii dif­ and thus provided invaluable service. His fers from U. sagittatum in lacking the no­ untimely death on April 27, 2002 prompts dose thickening of the spines and in hav­ us to name this new species to honor his ing a thickened platelet at the base of the devotion to aroids. spines. Despite some reports to the contrary it ACKNOWLEDGMENTS now appears that most of the stems of The authors are grateful for reviews of Zomicarpeae are rhizomatous. Although the manuscript made by Josef Bogner, Ju­ Filarum manserichense Nicolson was de­ lius Boos. For descriptions and micro­ scribed as being tuberous in the original graphs of pollen we wish to thank Valen­ publication it has proven to be rhizoma­ tina Tarasevitch, and also Michael Hesse tous, like Ulearum donburnsii Qulius and Heidemarie Halbritter from the Uni­ Boos, pers. comm.). The illustration and versity of Vienna for the micrographs of description of Filarum in Genera of Ara­ the pollen used in this manuscript. Credit ceae (Mayo et at., 1997) describe Fitarum is also given to Wilbert Hetterscheid for his manserichense as being tuberous, as does advice regarding relationships within the the original description by D. Nicolson Zomicarpeae, especially in regard to stem (Nicolson, 1966). On the other hand, Wil­ growth. bert Hetterscheid (Hetterscheid & Size­ more, 1997) redescribed the species as rhi­ LITERATURE CITED zomatous. Filarum, Ulearum and Zomi­ carpella have all proven to be rhizoma­ Bogner, J. 1997. New taxa of Araceae. tous. Only Zomicarpa appears to be Sendtnera 4:5-12. tuberous, based on the both the illustra­ Boyce, P. C. 1995. Plant portrait: Ulearum tions and description in the Genera of Ar­ sagittatum. Curtis's Bot. Mag. 12:135- aceae (Mayo et al., 1997). Though the rhi- 137. T.E. CROAT, B. FEUERSTEIN, 2002 43

Hetterscheid, W. 1. A. & M. Sizemore. The Genera ofAraceae. Royal Botanic 1997. Aroid profile, Filarum manser­ Gardens, Kew. 1-370. ichense Nicolson, Aroideana 20:57- Nicolson, D. H. 1966. Filarum, a new ge­ 59. nus of Peruvian Araceae. Brittonia 18: Mayo, S. J., J. Bogner & P. C. Boyce. 1997. 347-349.