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Rac EN Ys As Vat eas staat aii «bade : Mietatee) ry +f at at Si Tes eee hg oe Bet | Saki bi ba bd 8) a t a > be even, fh apenas upstate " mM An ; { ie j 7 ‘ Shae y's ’ hk y ir ee PAL Sat Toth (Ae ei ka ai a A Pe a eR HT ed | r %, wig Oak Le ae oy n nm) Mary yak pdb ree 4 3 2 3 : - * a | / ¢ ie * ‘ V . LIBRARY OF Tiik NEW YORK BOTANICAL GARDEN LIBRARY NEW YORK BOTANICAL GARDEN MEMOIRS OF THE NEW YORK BOTANICAL GARDEN VotumeE 10, Numser 1 The Botany of the Guayana Highland—Part III ; Bassetr Macutre AND JOHN J. WuRDACK AND CoLLaBoraTors = 1 Anatomy of Guayana Mutisieae. Part IJ. SHERWIN Cartguisr 157 Issued July 1, 1958 The Memoirs of The New York Botanical Garden are issued at irregular intervals in parts of various sizes. Approximately 500 pages will complete a volume. The subscription price of volume 10 is $10.00. Number 1 may be purchased separately for $3.50. Authors of papers may obtain separate copies of their contributions, printed at the same time as the issue, at cost price. For further information address the editor: H. W. Rickett The New York Botanical Garden New York 58, N. Y. tJ e we ° ° « J * @ 3 e Sees Moe MoviccMacdaciacMacMasiac! > ERRATA MEMOIRS OF THE NEW YORK BOTANICAL GARDEN Volume 10, Number 1 July 1, 1958 2: in table for style of Abolboda, for "alcerately" read "lacerately" 12, line 21: for "prevalance", read "prevalence" 17, line 8: for “aciularis", read "acicularis" 21, footnote: for "Rapaecaceae", read "Rapateaceae" 22, line 9 of key: for "terte", read "terete" 32, line 1: for "(US 1903798; Herb. Lehmannianum s.n. s. dat. K)", read "(US 1903798); Herb. Lehmannianum s.n. s. dat.(K)." Van line 21: for "Sipopa", read "Sipapo" line 35: for "peduncelo", read "pedunculo" line hl: for "gratatis", read "gradatis" 35, line 30: for "Schoenocephaleae", read "Schoenocephalieae" 36, line 17: for "bivaviatis", read "bivalvatis" line 36: for "aranged", read "arranged" line 37: for "savana", read "savanna" 37, first line of key: for "upply", read "upper" 39, 2nd line from bottom: for "les", read "less" “hO, line 1: for "Mayo", read "Moyo" line 41: for "subbassally", read "subbasally" 43, line 47: for "lagoona", read "laguna" line 49: for "dege", read "edge" 48, line 45: for "Parophytum", read "Potarophytum" 51, line 10: for "obvate", read "obovate" line 24: for "numtrous", read "numerous" 92, last line of key: for "3.5 mm", read "3.5 cm" 95, last line: for "6-", read "6-9" 97, line 4: for "Rhychanthera", read "Rhynchanthera" line 6: for "Rhyncanthera", read "Rhynchanthera" line 7: for "Khyncanthera", read "Khynchanthera" 98, line 2: for "aerolata", read "areolata'" line 12: for "cordiforia", read "cordifolia" 100, line 2: for "glabor", read "glaber" 101, line 38: for "7-11 3-3.5", read "7-11 X 3-3.5" 102, line 33: for "impersgicue", read "imperspicue" 106, line 27: for "Schules", read "Schultes" 108, line 22: for "2,6", read "2.6 mm" 109, line 47: for "subus", read "subtus" 115, line 2h: for "apise", read "apice" 120, line 19: for "denique", read "deinde" 125, line 17: for "Ttate", read "Tate" 129, line 9: for "Ttate", read "Tate" | 132, line 28: for "suggenus", read "subgenus" line 38: for "3-6.5 mm", read "346.5 cm! | afer wish Peon | Wel rie ot ate i cal 0 4 sal soot bale , | athe lietoe™: Beer, . ; er i mneaX x 3} r bao ny) Bie ea meton wt inoue nes ¢ ery ue Ted SOS. beer. Pid hs oe. cif Le ecu ey fic oe foe suse seat, as saci bic a ha ais a Side pionn: ‘baat : ih shh aes sear Cas {oe ds daa” | bauer at poet cena : eM i Megat -y ne ag eer 1 any 4 "i My Sieve ee, vi i Ph ee ia : iV % ah waft | ML \ i So Mini orl ooo Been I ok Mat glace 4 é c 8) or 4 a 7 a, ii a chi ay , ee ae It me ney: aT t) aa Om exe 30) , Were Goce tip 2 is y * aa: bd yon oP die tee we ii oe gl ii at: eels, iA ait reich uff ‘Biok j ‘, LY EEPE < rae rn y aay erg wy iret Ringe: ‘ ‘ stiles . roe at Ne RAIN lg wee Page #2. ERRATA | MEMOIRS OF THE NEW YORK BOTANICAL GARDEN Volume 10, Number 1 July 1, 1958 P, 133, line 33: for "mor", read "more" P, 134, line 2h: for "tingled", read "tinged" P, 135, line 6: after "Cano,", insert "2000 m alt., Jan 31, 1951, sparsely branched shrub to 1.5 m tall, flowers yellow with tube reddish basally. Cowan & ) Wurdack 31144, Serrania Paru, Rio Paru, Caro Asisa, Rio Ventuari, frequent, cumbre just south’ of head of valley of Camp Cano," line 15: delete line and insert "portion) is up to 1.4 cm long, the throat 1.7 cm long, the corolla- lobes 1 cm long," P, 137, line 5: for "rugolose", read "rugulose" line 37: for "acutifilia", read "acutifolia" P, 143, line 31: for "pattren", read "pattern" P, 144, line 32: for "collections, there", read "collections; there" P, 160, plate explanation: for "urdack 565", read "Steyermark & Wurdack 565" P, 162, line 33: for "intrepretation", read "interpretation" P, 164, line 15: for "is", read "are" P, 166, line 23: for "ontogny", read "ontogeny" P e FP . 168, line 8: for "palidsade", read "palisade" .» 169, last line of legend: for "portiton", read "portion" - 179, line 14: for "martianus", read "martiana" ne he bl eqn. ibe eT) Tae bf .* rm Tt eo. @ wd EG Re pom Lesbos wf - Pr iy Ae Le, es be Mi Pak out * 1 Poy Par 00 ve” fi a | + et ee Teer ais Tran pep } P p re is s FD Voriet if Bi hae sath tay, ves ney rato is ha ag ie te nh be x ; a of = ARS a me pad i ; 4, ABTA bh i tee aay Sh a Ns ei le Sed ae te AN Sp) My he wee NED este Se OTS Ts tee ewe ee Vol. 10, No. 1 - MEMOIRS OF THE NEW YORK BOTANICAL GARDEN July 1, 1958 THE BOTANY OF THE GUAYANA HIGHLAND—PART III BASSETT MAGUIRE AND JOHN J. WURDACK AND COLLABORATORS INTRODUCTION In Part III we continue to report on plant collections made in South Ameri- can Guayana. No additional field work has been done since the preparation of Part [I* of this series. However, at this writing we are making ready for immi- nent return to the region of the Casiquiare, and the Rios Pacimoni and Yatua in Amazonian Venezuela. There we are again to visit Cerro de la Neblina and the immediately outlying tabular mountains, all of the Roraima sedimentary complex. This, the nineteenth of our Guayana expeditions, is expected to be in the field for some six months. Field work and publication continue with the assistance of the National Science Foundation. Of the studies presented herein, those of the Xyridaceae, Rapateaceae, and Guttiferae have been made by Maguire, and of the Melastomataceae by Wurdack. Contributions of collaborators are as follows: Leguminosae and Rutaceae, Rich- ard S. Cowan; Myrtaceae, Rogers McVaugh; Combretaceae, B. Maguire and A. W. Exell; Apocynaceae, Joseph Monachino; and Bignoniaceae, N. Y. Sandwith. Sherwin Carlquist presents Part II of the ‘‘Anatomy of Guayana Mutisieae.’’ Illustrations have been prepared by Charles C. Clare, Alexis Khoury, Walter L. Graham, and Richard S. Cowan. XYRIDACEAE Traditionally, two genera have been recognized as constituting the small family Xyridaceae: Xyris with numerous species widely distributed in the Western Hemisphere, Europe, Africa, and Australia, and Abolboda with com- paratively few species confined to tropical South America (exclusive of Andea). Both have been recognized as homogeneous genera except for the anomalous Abolboda sceptrum, for which until recently there has been insufficient material for adequate diagnosis. Im Thurn on his Roraima expedition (1884-1885) collected the upper por- tion of a single flowering scape. Upon this specimen Oliver established the name Abolboda sceptrum.1 Later, McConnell and Quelch? (1894, 1898) succeeded in collecting only adult leaves and young plants. Apparently the plant was not again collected until Tate in 1928 obtained single specimens of flowering and fruiting plants from Cerro Duida.* Of these specimens, Malme, prominent stu- dent of the Xyridaceae who reviewed Tate’s material, wrote, in part, ‘‘forsitan novum genus sistat.’’ Subsequently, no fewer than 26 collections of A. sceptrum and its segregates have been made. Ample opportunity has been had to study the several popu- lations in the field. It has now become abundantly demonstrable that the plants known as A. sceptrum must be referred to a distinct genus, for which I propose the name Orectanthe (descriptive of the porrect flowers). On palynological evidence, Erdtman* has suggested that Abolboda and Xyris *The botany of the Guayana Highland—Part II. Mem. N. Y. Bot. Gard. 9: 235-392. 1957. 1 Trans. Linn. Soe. IT. Bot. 2: 286. 1887. 2 Trans. Linn Soe. II. Bot. 6: 69. 1901. 3 Bull. Torrey Club 58: 326. 1981. 4 Pollen morphology and taxonomy 453, 454. 1952. Oo 2 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN ‘ [vou. 10 , are familially inconsonant. Taxonomic evidence supports this suggestion.t Addi- tional morphologic data are now being sought further to elucidate the question. The following table indicates the more obvious contrastive and comparative features of the three genera which are at this time tentatively retained in the X yridaceae. Orectanthe Abolboda Xyris te. Habit. caulescent. acaulescent. acaulescent. 2. Leaf arrange- rosulate or cauline. basal, sometimes rosu- basal, often distichous ment. late, in at least one spe- and the sheaths strongly cies distichous and equi- equitant. tant. 3. Scape ebracteate. bracteate (commonly). ebracteate. 4. Sepals. Bie 2 (commonly). a 5. Corolla. irregular, yellow. regular, or somewhat ir- regular, yellow, rarely regular, blue, rarely white. 6. Staminodia. 7. Pollen grains. lacking. nonaperturate, spheroi- dal, spinose, 160—250 u in diameter. white. lacking, or simple and filiform. nonaperturate, spheroid- al, spinose, 75-150 w in diameter. bifureate, strongly peni- cillate. aperturate, sulecate, with- out spines. 8. Style. crateriform, penicillate, crateriform, penicillate, trifid above the middle, undivided. sometimes alcerately tri- the segments linear. fid. 9. Seed. strongly compressed, obliquely oblong, oblique- oblong, vertically striate. broadly winged, non- striate; mitten-shaped. Key to the Genera of the Xyridaceae 1. Sepals 3; flowers yellow. 9 ly striate. 2. Corollas irregular, porrectate, exceeding 5 em in length; staminodia lacking; seed strongly compressed, broadly winged; coarse caulescent herbs with rosulate or conferted cauline leaves. 9 1. Orectanthe. 2. Corollas regular, less than 3 em long; staminodia bifurcate, prominently penicil- late; seed oblong, vertically striate; acaulescent herbs with basal and often dis- tichoys but not rosulate leaves. 3. Xyris. 1. Sepals 2 (commonly); corollas blue, essentially regular, 3 em or less long; stamino- dia lacking or filiform and unbranched; seed obliquely oblong, spirally striate; acaulescent herb with basal, rosulate, or sometimes distichous leaves 2. 1. Orectanthe Maguire, gen. nov. Abolboda. Inflorescentia solitaria, terminalis capitata subglobosa, vel hemisphaerica am- plior, multiflora; floribus sessilibus porrectis singulis a bractea subtentis; brac- tearum cyclo vel duobus inferioribus sterilibus; floribus trimeris; sepalis 3, duobus lateralibus conduplicatis equitantibus valde carinatis faleate lanceolatis, tertio anteriore nec conduplicato nec carinato; corolla gamopetala aliquantum inaequaliter bilateraliter symmetricali porrecta, lobo posteriore maiore; stamini- bus 3, antheris adnatis linearibus 4-locularibus; granis pollinis maximis, 160—— +t Mr. A. A. Bullock has brought to our attention the formal and valid publication of the name of Abolbodaceae Nakai, in Ordines, familiae, tribi, [ete.] in praelectionibus anni 1937. 1958] THE BOTANY OF THE GUAYANA HIGHLAND—PART III 3 250 » diam., sphaeroideis, non aperturatis spinosis; ovario 3-loculari, placentis 6, axialibus 2—3-seriatis; stylo simplici ad basim a 3 valde recurvatis simplicibus appendicibus subtento; stigmate oblique terminali aliquantum fimbriato crateri- formi; seminibus valde compressis, late alatis, oblongis irregulariter mitriformi- bus; embryone recto. Herbae perennes caulescentes; laminis lineari-lanceolatis acute acutis, rosu- latis vel confertis. Typus: Orectanthe sceptrum (Oliver) Maguire. Orectanthe from 6gex70¢, stretched outward, av6o¢, flower; because of the porrectate flowers. Key to the Species of Orectanthe Plants essentially acaulescent, the stem 1 dm or less long; leaves densely rosulate; usu- ally unipedunculate; cephalar bracts commonly 3 in a single cycle. 1. Orectanthe sceptrum. Plants caulescent, the stems 4-15 dm long; leaves conferted but not rosulate; often pluripedunculate; cephalar bracts commonly 6 in 2 eyeles. 2. Orectanthe ptaritepuiana. 1. Orectanthe sceptrum (Oliver) Maguire, comb. nov. Figure 1. Caudex short ; leaves numerous, stiffly rosulate, linear-lanceolate (1.2) 1.5-4.5 em wide, (10) 15-30 em long, conspicuously brown-margined, the base broad and more or less clasping, the apex pungently acute or acuminate; scapes terete, usually solitary, 3-12 dm high, subtended at the base by 2 or 3 lanceolate bracts 2-8 em long; heads ovate before anthesis, enclosed by 3 (2) ovate to lanceolate sterile bracts, at maturity globose-hemispheric, the flowers porrect, each sub- tended by an ovate-lanceolate bract shorter than the sepals; sepals 3, the lateral keeled pair 5-7 cm long, the anterior sepal not keeled and somewhat shorter ; corolla yellow (5) 6-8 em long, the tube + equaling the lobes; style shorter than the corolla; capsule obovate-obpyriform, 3-celled, loculicidal, 1.5-2.0 em long; seed compressed, winged, orbicular or oblong, irregular, mitten-shaped, (3.5) 4-5 mm long. Key to the Subspecies of Orectanthe sceptrum Subtending cephalar bracts lanceolate, 3-5 em Jong; herbage glaucous. subsp. sceptrum. Subtending cephalar bracts ovate, 1.5—-2.0 em long; herbage not glaucous. — subsp. occidentalis. Orectanthe sceptrum subsp. sceptrum. Abolboda sceptrum Oliver, Trans. Linn. Soe. II. Bot. 2: 286. 1887. Plants ordinarily 1.0-1.5 m high; herbage glaucous; cephalar bracts lanceo- late to ovate-lanceolate, 3-5 cm long, very acute or merely acutish. Type: ‘‘Roraima summit,’’ im Thurn 312 (K). The type consists wholly of a Single ‘‘. . . well developed capitulum and 8~9 inches of its scape.’’ Distribution: plants of open marshy or boggy areas, and wet savannas, Brit- ish Guiana Pakaraima Mountains and the Gran Sabana, Venezuela, at 500 m upwards to cumbre summits at more than 2000 m altitude. Specimens examined: BRITISH GUIANA: Pakaraima Mountains: perennial herb with solitary scape and basal rosette of linear-lanceolate, brown-margined leaves, occasional in marshy area, Imbaimadai Savannas, Upper Mazaruni River, at 550 m altitude, Oct 24, 1951, Maguire §& Fanshawe 32254; acaulescent marsh herb, scape 1 m high, flowers pale yellow, savannas 15 km. n. Mt. Ayanganna, at 1200 m altitude, Feb. 2, 1955, Maguire, Bagshaw & Maguire 40567. VENEZUELA: Gran Sabana, Edo. Bolivar: ‘‘Roraima summit,’’ £. F. im Thurn 312 (holotype, K); Emerald Swamp, southwest-facing slope at 1520 m alt., Sept 25, 1944, Steyermark 58615 (NY, F); summit Mt. Roraima at 2620-2740 m alt., Sept 27, 1944, Steyermark 58861 (F); frequent, marshy savanna, between San Rafael and Enemasiec, Fig. 1. a-n, Orectanthe sceptrum. a, habit, < 1/,; b, inflorescence, X 14; ¢, flower in natural posture, K %; d, corolla spread out, X 14; e, sepals, anterior view showing subtend- ing bract, posterior view, X%4; f, diagrammatic representation of cross-section of keeled lateral — sepals, and concave inner sepal, X 1; g, cross diagram of flower; h, ventral view, * 2, and cross section, X 4, of anther; i, pistil, X 14; j, k, 1, stages in developing stigma, « 4; m, face view, and n, cross section of seed, & 5; drawn from Magwire 33189. i) | 1958 | THE BOTANY OF THE GUAYANA HIGHLAND—PART III ca. 12 km w. Ilu-tepui at 1200 m alt., Feb 15, 1952, Magwire 31180; marshy places in open savanna, Mesa Grande, Ilu-tepui, at 1600 m alt., Mar 12, 1952, Maguire 33342; Ptari-tepui, sw-facing shoulder, Nov 2, 1944. Steyermark 59787 (F) ; Ptari-tepui at 1615 m alt., Nov 15-17, 1944, Steyermark 60244 (F); Kavanayén, May 28, 1956, Lasser 1826 (NY); Kavanayén at 1300 m alt., Mar 30, 1952, Maguire 33680. Orectanthe sceptrum subsp. occidentalis Maguire, subsp. nov. Plantae vulgo 5-10 dm altae; gemmis axillaribus frequentibus; non glaucis; bracteis cephalaribus ovatis vel lanceolato-ovatis, 1.5—-2.0 em longis, acutiusculis. Type: scapose perennial herb 1 m high, scapes reddish, frequent in marshy scrub savanna, Southeast Escarpment at 1900 m alt., Cerro Huachamacari, Terr. Amazonas, Venezuela, December 11, 1950, Maguire, Cowan & Wurdack 30140 (NY). Paratypes: VENEZUELA: Amazonas: Cerro Sipapo: wet ledges, West Peak at 1800 m, Dee 20, 1948, Maguire § Politi 27777; savanna bog, Lower Cano Negro, at 1400 m alt., Jan 1, 1949, Maguire & Politi 28106 ; Upper East Basin at 1900 m, Jan 14, 1949, Maguire & Politi 28451; marshy area, North Mountain at 1500 m alt., Jan 25, 1949, Maguire 5: Politi 28569 ; savanna, South Escarpment at 2100 m alt., Jan 26-28, 1949, Maguire, Politi & Magwre 28645 ; savanna, South Basin at 2000 m alt., Jan 26-28, 1949, Maguire, Politi & Maguire 28678. Summit Mt. Duida: at 6300 ft. alt., Nov 26, 1928, Tate 398. Cerro Huachamacari: savanna at 1700 m alt., Dee 4, 1950, Maguire, Cowan §& Wurdack 29822; summit East Ridge at 1820 m alt., Dee 8, 1950, Maguire, Cowan §& Wurdack 30039; Escarpment Ridge 1800 m alt., Dee 14, 1950, Maguire, Cowan §& Wurdack 30213. West Escarpment, Cerro de la Neblina at 1750 m alt., Jan 15, 1954, Maguire, Wurdack §& Bunting. 37303. Distribution: marshy areas and wet savannas, cumbres of the sandstone massifs of Venezuelan Amazonas. Specimens from Cerro Sipapo are usually smaller than those from elsewhere and commonly develop offshoots from some of the leaf-axils, and thus seem to represent a distinct minor variant. 2. Orectanthe ptaritepuiana (Steyermark) Maguire, comb. nov. Abolboda ptaritepuiana Steyermark, Fieldiana Bot. 28: 104. 1951. Caulescent perennial herb; herbage not glaucous; stems 0.5—2.0 m tall; leaves numerous, rather laxly conferted but not rosulate, linear-lanceolate, obviously or not at all brown-margined, pungently acute, 15-30 em long, 1-5 em broad; peduncles more often several (solitary in the type): sterile bracts in 2 whorls of 3’s; inflorescence and flowers otherwise similar to those of O. sceptrum. Type: in dense thickets and Bonnetia roraimae forest southwest facing shoul- der, Ptari-tepui, Edo. Bolivar, Venezuela, Nov 2, 1944, Steyermark 59760. Distribution: at higher elevations, 1500-2000 m alt., on Roraima Sediments, Edo. Bolivar, Venezuela. Specimens examined: VENEZUELA: Bolivar: Ptari-tepui, Steyermark 59760 (holotype F, 2 sheets). Cerro Guaiquinima: coarse perennial herb dying on flowering; bracts 4-5, open savannas 1 km south of Cumbre Camp at 1800 m alt., Dee 29, 1951, Maguire 32823; young plants with a single scape, mature plants with 3-6 scapes, primary cephalar bracts of two. whorls of 3’s, marshes and Brocchinia thickets 1 km e of Cumbre Camp at 1800 m, Maguwire 32786 ; marsh-bog herb, peduncles as many as 10, North Valley at 16-1700 m alt., Jan 4, 1952, Magwire 32979; valley below se Escarpment at 16-1700 m alt., Jan 7, 1952, Maguire 33022; marsh-bog, sometimes with branched stem, peduncles (1) 3-10, Cumbre Camp at 1800 m alt., Jan 9, 1952, Maguire 33035. Gran Sabana: perennial herb, stems 1 m high, leafy, scapes -5, heads subnectant, scrub type associated with Brocchinia at 2400-2500 m alt., Ilu-tepui, March 17, 1952, Maguire 33467; herbaceous perennial, abundant in large colonies, escarpment face and talus, Sarvén-tepui, Chimantaé-tepui, at 1900-2050 m alt., Jan 13, 1953, Wurdack 34134-A ; herbaceous with elongated erect leafy stems up to 10 feet tall, elfin formation ApAcara-tepui, Chimanta-tepui, June 19, 1953, Steyermark 75698 (F, 3 sheets, NY); stems erect, 3-8 ft. tall, Apacara-tepui, Steyermark 75885 (F, 2 sheets, NY, 2 sheets); herb with leafy stem, scapes 1—5, common, bogs at 2200 m alt., Serra do Sol (Uei-tepui), Maguwire §& Maguire 40413; Chimanté-tepui, 1955, Steyermark and Wurdack 523, 905, 1221. \ 6 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Orectanthe ptaritepwiana is distinguished from O. sceptrum largely by means of vegetative differences, and to some extent on geographical grounds. The first is characterized by well developed leafy stems and the prevalence of 3 or more peduncles, as contrasted to O. sceptrum which has very short stems and densely rosulate leaves. Although variable in size and trivial detail, there seems to be no way conclusively to separate the two species by floral distinction. They both occupy the geographical range in eastern Guayana, but only O. sceptrwm is to be found in western Venezuelan Guayana. Both species inhabit upper cumbre habitats; only O. sceptrwm occurs also at low altitude (1000 m) on the Gran Sabana. 2. Abolboda Kunth in Humb. & Bonpl. Pl. Aequin. 2: 25, 109. pl. 114. 1809. Key to the Species of Abolboda 1. Seapes provided with 1 or more pairs of subopposite bracts (exclusive of basally subtending members. 2. Seapes commonly with 2 (occasionally 3 or 4) pairs of bracts, or sometimes with only one pair, then exceeding 3 dm in height; herbs normally 3 dm or more high. 3. Spikes simple, not glomerate. ; 4. Seapes simple, with a single terminal spike. 5. Spikes globose or ovoid, the paired sepals and subtending bracts outwardly arched or divergent at maturity, bracts % or less as long as the sepals. 1. A. grandis. 5. Spikes eylindric or elliptic-cylindric; the paired and subtending bracts ascending or strict at maturity; bracts more or less equaling, or at least more than 14 the length of the sepals. 2. A. macrostachya. 4. Seapes branched, or if simple, with 1 or 2 axillary spikes below the terminal. 6. Sceapes paniculate; spikelets 2—4-flowered, solitary and terminal on each branch; leaves numerous, densely rosulate, gramineous, 5-10 mm long, 1.5-3.0 mm broad, narrowly acute, mucronate. 3. A. paniculata. 6. Scapes simple; spikelets 1-flowered, terminal, and in addition axillary spike- lets from the penultimate or antipenultimate vaginal bracts; leaves numer- ous, densely rosulate, narrowly linear, triquetrous, obtuse, abruptly mu- cronulate, 2-3 em long, ca. 1 mm broad. 4. A. uniflora. 3. Spikes glomerate; Rio Guainia drainage, Venezuela (and Colombia). 5. A. glomerata. 2. Scapes commonly with a single pair of bracts. 7. Spikes cylindric or subeylindric, exceeding 2 cm in length; leaves linear, obtuse, 20-30 em long, considerably exceeding % the length of the scapes; upper Rio Orinoco and Rio Guainia drainages. 6. A. linearifolia. 7. Spikes ovate or elliptic, 1.5 em or less long. 8. Leaves commonly % or more the length of the scapes. 9. Bracts immediately subtending and much exceeding the spike; spikes ovate, 1.0-1.5 em long; sepals and subtending bracts ascending or strict; leaves pseudo-equitant, gramineous, acute, 10-30 em long, more or less equaling the scapes. 7... A; spruce. 9. Bracts not immediately subtending the heads, affixed at more or less the middle of the scape; flowers and subtending bracts arcuate or subdivaricate. 10. Bracts of the scape narrowly lanceolate, attenuately acuminate, 2.0—2.5 em long; cephalar bracts strongly productate; sepals 16-20 mm long; plants 10-15 em high; Yapacana Savannas, Amazonas, Venezuela. 8. A. bella. 10. Braets of scapes lanceolate, abruptly acute, mucronulate, 1.0-1.2 em long; cephalar bracts ovate-lanceolate, ca. 7-8 mm long, short-produe- tate; sepals 9-10 mm long; plants 6-8 em high; campos do Ariramba, Rio Trombetas, Para, Brazil. 9. A. abbreviata. 8. Leaves commonly 14 or less the length of the scape. 11. Leaves plane, or more than 1 mm broad. 12. Leaves. entire; bracts of the scape 1.0-3.5 em long. 13. Flowers 5 or more per spike; bracts of the scape 2.5-3.5 em long, the 1958 | THE BOTANY OF THE GUAYANA HIGHLAND—PART III 7 first cephalar bracts 7-9 mm long, acute, mucronulate; campos and chapadas, Matto Grosso, Minas Gerais, Goyaz, southern Para, Brazil. 10. A. poarchon. 13. Flowers commonly 2—4 per spike; bracts of the scape 1.0-1.5 (2.0) em long, the first cephalar bracts 8-12 mm long, acutish, bluntly short- productate; savannas and campos, range of the preceeding, and Ama- zonas, Brazil; Vaupés, Colombia; and Amazonas and Bolivar, Vene- zuela. 11. A. pulchella. “12. Leaves serrulate-ciliate; bracts of the scapes 4.0 em long; cumbre of Cerro Neblina, Amazonas, Venezuela. 13. t hig. 11. dissitiflora. h, habit, * 0.5; l-m, large and small stamens, * 1.5. MEMOIRS OF THE NEW YORK BOTANICAL GARDEN Z . —S y7 . ns (= UW. CS ciara Gn Sa OA ; 50000 0) OU... a—g, Comolia serpyllacea. sO hed, pistil,. x 2.04 -e-1. large and small stamens, X 3; g, leaf, 5. h-m, Tibouchina [vou. 10 , DRESS FRB _ AS x 23s AM Gey CINE SES AS =s N28 at Bae ) ESS NNR ESSN Bey Aa BSZ5e WY UE) YN, SZSZ8 Bi he 8) Dw) gS f Why Cs 99) -'SS — 7 5 J “) J SSS . Gav ¥ Vy ZSZs K ASO y? y wd SSS Ay TE ami eee SESE : Yu wy a2 ZS § Y iy Jo Wouy gS § BASED RV RY =ZSZa2 NV: ARC ei SCNSSN SN zi Val ey Dal ys) : SASZ SNZie aN Gis Daher SANZ SNZA y Ve Sr deg aly PSN WN Ar elec) Way) OS = S ah M ck V J Voy GES= S : ¥ Mee in SS SNZA VAR Gd Ones Lo Se SNZE Suey ddr eh SINZ a gee al Ae peg i A SS 4 ARLES yf SOT SS pes ¥ YAP) Sass SW YULULY 7 J Aids =! re Y N ) way = Ii Y, LBP Bae PRC eho) Ws % yD Bd ts Ea oe > 2 = > > 2 ee = > —~ < a, habit, X 1; b, petal, X 2; ¢, hypanthium and calyx, 1, petal, p< leas a5 hypanthium and calyx, X 1.5; k, pistil, X 1.5; n-s, upper and lower leaf surfaces of Tibouchina spp. 1958 | THE BOTANY OF THE GUAYANA HIGHLAND—PART III 105 eciliate. Stamens slightly to markedly anisomorphic, the larger antesepaler. Anthers linear-subulate, arcuate, 1-pored; connective prolonged below the thecae into a bilobed ventral appendage, with a dorsal caudal appendage on the con- nective between the thecae and filament insertion. Ovary free, 4-locular, multi- ovulate, glabrous or apically puberulous or setose; style thin, long-exserted ; stigma punctiform. Fruit capsular, the hypanthium smooth or 8-costate; capsule 4-locular ; seeds cochleate, strongly muriculate. Generotype: P. coriaceum DC. Key to the Species of Pachyloma Leaf-surfaces glabrous; apex of the ovary glabrous (rarely sparsely puberulous). Leaf-margins strongly incrassate; leaves notably 5-nerved. 1. P. coriaceum. Leaf-margins not thickened; leaves mostly 3-nerved, occasionally faintly 5-nerved. 2. P.huberioides. Leaf-surfaces conspicuously glandular-setulose; apex of the ovary densely coronate- setulose with long glanduliferous hairs. 3. P. setosum, 1. Pachyloma coriaceum DC. Prodr. 3: 123. 1828. P. coriaccum var. a glaberrima DC. Prodr. 3: 123. 1828. P, coriaceum var. 8 subsetosum DC. Prodr. 3: 123. 1828. Rhexia pachyloma Schr. & Mart. ex DC. Prodr. 3: 123. 1828. Nomen. Rhexia bicuspis Schr. & Mart. ex DC. Prodr. 3: 123. 1828. Nomen. Rhexia amazonum Schr. & Mart. ex DC. Prodr. 3: 123. 1828. Nomen. Heteronoma pachyloma Mart. Noy. Gen. Sp. 3: 140. pl. 273. 1829. Shrub, the terete to obscurely quadrangular branches glabrous. Petioles 3-10 mm, glabrous; leaf-blade 5-10 «& 2-5.5 em, oval to ovate-oval, the apex short- acuminate, the base rounded to subcordate, the margins notably inecrassate and recurved, entire or evanescently serrulate, 5-nerved, the nerves narrowly im- pressed above and prominulous-incrassate below, the secondary nervation nar- row and rather obscure, obscurely punctate beneath at maturity, glabrous. Pedicels 0-12 mm below the bracteoles, 1-3 mm above, the early-caducous bract- eoles oblanceolate and 1-3 & 0.5-1 mm. Hypanthium (6—) 8-11 mm long, long- campanulate, the fleshy deltoid to rounded calyx-lobes 0.5-1 mm long. Petals 15-20 & 8-9 mm, ovate to elliptic, short-acuminate, seto-mucronulate. Stamens glabrous, the filaments 7.5-11.5 mm lone antesepalous stamens with thecae 10-12 mm long, the connective prolonged 0.6—3.5 mm to the filament insertion, the ven- tral acute spurs 0.2-0.5 mm long, the dorsal seta 1.8—2.2 mm lone’; alternisepal- ous stamens with thecae 9.5-11.5 mm long, the connective prolonged 0.4—0.7 mm to the filament insertion, the ventral acute spurs 0.6-1 mm long, the dorsal seta 1-1.4 mm long. Style 16-17 mm long, glabrous or sparsely glandular- setulose; ovary glabrous. Type Collections: Martius s. n. ‘‘Prov. R. Negro’’ (M, holotype of P. coriaceum and Rhexia pachyloma) ; Martius s. n. ‘‘Prov. Para’’ (M, holotype of P. coriaceum var. glaber- rima and Rhexia bicuspis); Martius s. n. ‘‘Ararocoara’’ (M, holotype of P. coriacewm var. subsetosa and Rhexia amazonum). n. T. spruceana (Maguire, Wurdack, & Bunting 36136), K 1; 0, T. aspera (Maguire & Maguire 40461), X 1; p, T. fraterna (Magwire § Fanshawe 32508), X 1; q, T. dissitiflora (Maguire, Wurdack, §.Bunting 37053), X 1; r, T. duidae (Cowan & Wurdack 31202), *K 1; 8, T. sipapoana (Maguire § Politi 28466), XK 2. % 106 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Additional Material: COLOMBIA: Vaupés: Cerro Yapoboda, Rio Kuduyari, Schultes & Cabrera 14249 (NY, US); Cerro Kafiendé, Rio Kubiyt, Schultes § Cabrera 18339 (NY, US), 183889 (US); Mesa de Yambi, Rio Karuru, Schultes g& Cabrera 19127 (NY, US); R-o Parana Pichuna, Schultes ¢ Cabrera 19927 (NY); Rio Vaupés, Mitu, Humbert 27324 (P). The Munich specimens apparently were never annotated by Cogniaux or Triana. None of the sheets bear de Candolle annotations but perhaps were seen by him before being mounted. All three Munich sheets bear Schrank diagnosis slips, although the diagnosis attached to the ‘‘Rhexia bicuspis’’ sheet is for Leandra quinquenodis (DC.) Cogn. Drs Baehni and Weibel supplied the infor- mation that there is only one herbarium sheet of Pachyloma in the de Candolle herbarium, with a packet containing leaves and a flower along with a label written by A. P. de Candolle “‘Rh. pachyloma L. Bresil M. Martius 1827’’; two other labels, not attached to any plants and not in de Candolle’s script, bear brief French diagnoses of ““R. amazonum a ovatae’’ and ‘‘R. amazonum B lanci- folia.’’ Baehni and Weibel also supplied the following information: Martius kept up a close correspondence with A.-P. de Candolle and called him ‘‘my dear friend.’’ In one of his letters to DC., dated March 13, 1827, he wrote; ‘‘Je viens de vous envoyer par la messagerie le second cahier du 2° volume de mes Nova Genera.’’ There is one thing of great interest in the Memoires et souvenirs de Augustin-Pyramus de Candolle (issued in 1862). De Candolle says that he went to Munich in Fall 1827 to attend a meeting of the German men of sciences; he was received there with great kindness by M. de Martius; he writes (p. 377) : ‘*Pendant les quelques jours que j’y ai passés, j’avais vu les portions de Vherbier de Martius relatives aux familles dont je m’occupais pour le troisieme volume du Prodromus.”’ The single-branch stamen dimorphism in the Schules & Cabrera collections is marked. Markedly dimorphic antesepalous and alternisepalous stamens are found on: 74249. (NY), 18339 (NY), 18389 (US), and: 19127 (NY jee euniso- morphism is seen in 14249 (US) and 18339 (US). The NY sheet of 19927 has one isomorphic branch and a separate anisomorphie branch. An unmounted US sheet of 18339 has dimorphic stamens but nearly glabrous hypanthium and glabrous style; the NY sheet of this number has moderately glandular hypanthia and glandular styles. The hypanthial and pedicellar glandulosity is so variable that there seems to be no reason for maintaining De Candolle’s varietal distinc- tions. On two of the Martius sheets at Munich, long connective prolongations are visiblé; the sheet of P. coriaceum var. glaberrimum now has no intact stamens. 2. Pachyloma huberioides (Naud.) Triana, Trans. Linn. Soc. 28: 64. 1871. Urodesmium huberioides Naud. Amn. Sci. Nat. ITI. 15: 338. 1851. Pachyloma scandens Ducke, Arch. Inst. Biol. Veg. Rio 2: 65. 1935. Weak shrub 0.5-2.5 m, the terete branches glabrous. Petioles 2-8 mm, gla- brous; leaf-blade 4-10 « (0.6—) 1-5 cm, lanceolate to ovate-elliptic, the apex acuminate, the base rounded to subecordate, the margins obscurely to notably. denticulate when young (obsctirely so with age) and not incrassate (the blade otherwise glabrous), 3-nerved or weakly 5-nerved, the nerves impressed above and prominulous below, the secondary nervation obscure to finely prominulous, sparsely punctate beneath at maturity. Pedicels 0.6-12 mm long below the bracteoles, 1-3 mm above, the caducous linear to oblanceolate bracteoles glabrous or sparsely glandular-setulose. Hypanthium 6-12 mm long, suburceolate to long- 1958 | THE BOTANY OF THE GUAYANA HIGHLAND—PART III 107 campanulate, glabrous to sparsely glandular-setulose, the fleshy deltoid to round- ed calyx-lobes about 1 mm long and usually sparsely glandular-setulose. Petals 15-20: 8.5-9.5 mm, ovate to ovate-oval, acute, mucronulate. Stamens usually olabrous, the filaments 7.5-13 mm long; antesepalous stamens with thecae 8.5-13 mm long, the connective prolonged 1—3.5 mm to the filament insertion, the ventral acute spurs 0.4-1 mm long, the dorsal seta 2.5-7 mm long; alternisepalous stamens with thecae 9-11.5 mm lone, the connective prolonged 0.5-0.9 mm to the filament insertion, the ventral acute spurs 1-2 mm long, the dorsal seta 2.5-4.5 mm lone. Style 15-27 mm lone, glabrous; ovary glabrous or sparsely elandular-puberulous. Seeds 0.9 & 0.6 mm, the tubercles about 0.06 mm high. Type Collection: VENEZUELA: Rio Atabapo near San Baltazar, Bonpland 936 (Holo- type and isotype, P). Additional Material: VENEZUELA: Amazonas: Catio Cupaven, Rio Orinoco near mouth of Rio Atabapo, Level 68 (NY); Yapacana savannas, Magwire, Cowan, § Wurdack 30579 (NY), Maguire, Wurdack, & Bunting 36609 (NY); Sabana Grande, Esmeralda, Maguire & Wurdack 34690 (NY); Caio Pimichin sabanita near Pimichin, Maguire, Wurdack, ¢ Bunting 36390 (NY); savanna on right bank of Rio Pacimoni 50 km above mouth, Maguire, Wuwrdack, § Bunting 37558 (NY). BRAZIL: Amazonas: lower Rio Curicuriary, Ducke H.J.B.R. 24105 (isotypes of P. scandens, P, US); Rio Negro, Uanauaea, Froes 223829 (IAN, NY, U). The collections 36609 and 36390 are fruiting only; I have been unable to find any ovarial pubescence in collections other than these two, even in one available fruiting branch of 34690. Of all the recent collections, Level 68 is the best match for the holotype. In part of 34690, the filaments apically have a few glandular hairs. Flower size varies greatly within the cited specimens. Ducke 24105, MWB 37558, and Froes 22329 have approximately isomorphic stamens. MW 34690 has quite anisomorphic stamens, while Level 68 has considerable variability (with generally anisomorphie stamens). 3. Pachyloma.setosum Wurdack, sp. nov. A congeneribus differt foliorum superficie glanduloso-setosa, ovaril apice setoso-coronato setis glanduliferis. Frutex 0.2-2 m; rami congesti teretes cum petiolis pedunculis pedicellisque modice persistenterque glanduloso-setosi. Petioli 2-4 mm; lamina 3-8.5 1.5- 3.9 em, ovalis apice rotundata mucronulata basi rotundata trinervia vel nervis duobus obsoletis inframarginalibus additiciis; nervis primariis supra anguste impressis subtus prominentibus, nervis secundariis inconspicue evolutis, supra sparse conspicueque glanduloso-setosa setis ad 4 mm longis glandulis caducis, subtus sparse glanduloso-setosa praecipue ad nervos superficie cum glandulis sessilibus 4-6/mm? ornata, marginibus glanduloso-setosis. Inflorescentia termi- nalia e dichasiis paucis composita; pedicelli 1-13 mm infra bracteolas, 1-3 mm supra; bracteolae caducae oblanceolatae vel lineares 1.1-3 & 0.5-1.5 mm, glandu- loso-ciliolatae. Hypanthium ecylindrico-campanulatum 9-10 mm longum con- spicue glanduloso-setosum calycis lobis obtusis inconspicuis. Petala 16.5-17 « 11.5-12 mm, ovalia apice breviter cuspidata extus apicem versus persparse elanduloso-setosa. Stamina isomorphica; thecae 10-11 mm lonegae, connectivo sub loculis 1 mm protracto appendicibus ventralibus acutis 1.5 mm longis, seta dorsali 3.5-4 mm longa, paulo supra insertionem filamenti inserta, filamento 9 mm longo glabro. Stylus 16-17 mm longus. glaber; ovarium 6.5 mm altum, apice setis multis glanduliferis coronatum; hypanthium fructiferum 15 « 10 mm obtuse 8—costatum; semina 1 & 0.9 mm cochleata valde muriculata. Type: sabanita 1 km west of Maroa, elev. 130 m, Rio Guainia, Terr. Ama- - a 108 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 zonas, Venezuela, Nov 25, 1953, Bassett Maguire, John J. Wurdack, & George 8S. Bunting 36397 (NY). Paratypes: same locality, frequent shrub, petals magenta, filaments white except for pink stripe just below anthers, April 17, 1953, Ma- guire & Wurdack 35717. Adelobotrys fruticosa Wurdack, sp. nov. A. barbatae Tr. affinis sed foliis plerumque minoribus foliorum indumento subtus densissimo appresso inflorescentia non pedunculata calycis dentibus maioribus. Rami juveniles dense rufo-villosi demum cinerei glabrati. Petioli lati, 3-10 mm longi; lamina 4-11 * 2-4.5 em, elliptica apice acuta basi obtusa vel trun- cata J—-7-nervia nervis primariis supra et subtus leviter elevatis nervis secun- dariis supra paulo elevatis subtus obscuris, supra primum villosula demum glabrata, subtus densissime strigulosa setis gracillimus primum rufescentibus demum cinereis tarde in lamellis irregularibus dehiscentibus; folia demum deflexa. Inflorescentia capitata sessilis vel maxime ad 1 cm pedunculata in toto cum pedicellis hypanthiis calycibusque densissime rufo-strigosa pilis gracilibus. Flores 5-meri breviter (ad 5 mm.) pedicellati, alabastris submaturis solum notatis. Hypanthium 6 & 4 mm; calyx ad basim 1 mm coalitus, lobis ovatis 2.5 mm longis, dente exteriore 1.2 mm longo acuto. Petala late obovata obtusa. Stamina dimorphica, maioribus cum antheris 4.7 mm longis, connectivi basi acuta 0.8 mm producta appendice dorsali adscendenti 2.5 mm longa, apice breviter emarginata, minoribus cum antheris 2.6 longis, connectivi basi acuta 0.6 mm producta anpen- dice dorsali adscendenti 2.5 mm longa apice rotundata. Ovarium 3-loculare elabrum apice truncatum. Type: shrub 0.3-1.5 m, frequent in sabanita on right bank of Cano Pimichin 1 km above Pimichin, elev. 140 m, Rio Guainia, Terr. Amazonas, Venezuela, Noy 24, 1953, Bassett Maguire, John J. Wurdack, & George S. Bunting 36392 (NY). Paratype: infrequent in woodland near Savanna No. 1, northwest base of Cerro Yapacana, elev. 125 m, Rio Orinoco, Terr. Amazonas, Venezuela, Nov 20, 1953, Maguire, Wurdack, & Bunting 36608. A. barbata has leaves with petioles 1.5-3 em long and blades up to 20 * 8 em; the lower leaf-surfaces are moderately covered with stout semi-erect mal- pighian hairs; the inflorescence is a short but definite panicle and the calvx proper is only indistinctly lobed. A. barbata is known from Maroa. Venezuela, and the ypper Rio Negro drainage (Rios Vaupes, Piraparana, and Icana). Adelobotrys duidae (Gleason) Wurdack, comb. nov. Meriania duidae Gleason, Bull. Torrey Club 58: 419. 1931. The umbellate inflorescence and fine brown cauline pubescence make this species much more congruous with Adelobotrys, where the nearest relatives, A. fructicosa Wurdack and A.. barbata Tr., are also both shrubs in a predomi- nantly vining genus. The persistent pubescence of the upper leaf-surface and the non-projecting exterior calyx-teeth immediately distinguish A. duidae; the species is known only from the cumbres of Cerros Duida and Huachamacari. — Graffenrieda hitchcockii Gleason var. parvifolia Wurdack, var. nov. A var. hitchcockii differt foliis minoribus ovatioribus subtus subglabratis. Paniculae angustiores cum fioribus paucioribus. Type: shrub or small tree 3 m high, petals white, stamens yellow, occasional 1958 | THE BOTANY OF THE GUAYANA HIGHLAND—PART III 109 near Camp Yutaje, elev. 1300 m, Cerro Yutaje, Terr. Amazonas, Venezuela, Feb 6, 1953, Bassett Maguire & Celia K. Magwire 35073 (NY). Paratypes (all from Terr. Amazonas, VENEZUELA): Cerro Yutaje, Maguire & Maguire 39091, 35232; Cerro Coro-Coro, Maguire §& Maguwire 35489; Cerro Guanay, Maguire, Phelps, Hitchcock, §& Budowski 31730. Var. hitchcocku has nearly elliptic leaf-blades mostly 12-25 & 9-13 em with rather persistent reddish puberulence; the floriferous parts of the panicles are 10-21 & 10-15 em. It is represented, apart from the type collection, by Maguire & Maguire 35338 and 35360 (both from Yutaje). Var. parvifolia has ovate- elliptic leaf blades 7-10 (-14) & 3-6 (-9) em with acute to acutely rounded apices ; the pubescence of the lower leaf-surface is chiefly confined to the nerves; and the flowering panicles are 7-13 & 5-7 em, expanding up to 20 * 9 em in fruit. The flowers of both varieties are either 4- or 5-merous; the ovary is pre- dominantly 3-celled, rather than 4-celled as originally described. Graffenrieda rufa Wurdack, sp. nov. G. hitcheockii Gl. remote affinis sed foliis minoribus acuminatis. Ramulorum novellorum petiolorum foliorum novellorum alabastrorumque pu- bescentia rufescentia densa granulosa. Petioli 8-15 mm; lamina integra 3.5-6.5 x 2-4 em, ovata apice breviter gradatimque acuminata basi paulo cordata D-nervia, nervis secundariis supra inconspicue impressis subtus paulo elevatis, supra et subtus primum dense granuloso-puberula, supra demum glabrata et nitidula, subtus demum persistenter dense subtiliterque cinereo-furfuracea. In- florescentia terminalis pluriflora; alabastra elliptica apice hebetiacuta 5-mera. Staminum connectivum postice cum appendice acuta antice exappendiculatum. Ovarium 3-loculare, dense granulosum apice in collum breve protractum. Fructus 1-2 mm pedicellatus ; fructuum sepala (vel dentes exteriores ?) ca. 2 mm longa, acuta; hypanthium fructiferum 44.5 & 3.5 mm; semina pyramidata laevia 0.7-0.8 & 0.3 mm. Type: shrub to 4+ m, in bud, rare in cumbre between between Summit Camp and southwest escarpment, elev. 1850 m, Cerro Huachamacari, Terr. Amazonas, Venezuela, Dee 17, 1950, Bassett Maguire, R. S. Cowan, & J. J. Wurdack 30282 (NY). Paratypes (all Terr. Amazonas, VENEZUELA): shrub or small tree to 5 m, inflores- cence tawny, frequent in cumbre of Cerro Huachamaeari, elev. 1700 m, Dee 4, 1950, Maguire, Cowan, & Wurdack 29817 ; shrub 5-8 ft tall, leaves dark green above, rufous brown below and on petioles and young stems, moist exposed cliff face at 1600 m elev., southeastern-facing sandstone bluffs near Cano Negro (tributary of Cafio Iguapo), Cerro Duida, Aug 26, 1944, J. A. Steyermark 58044, Graffenrieda fruticosa Wurdack, sp. nov. G. rufae Wurdack remote affinis, sed foliis minoribus triplinerviis non acuminatis venis secundariis subtus vix evolutis, inflorescentia vix evoluta. Ramuli novelli cum petiolis foliis subtus hypanthio calyceque densissinie -pallideque rufo-fulvi. Petioli 4-9 mm; lamina integra 10-25 & 6-10 mm (in ramis sterilibus ad 40 & 20 mm) elliptica vel paulo ovato-elliptica apice hebetiacuta basi late acuta breviter (0.5-1 mm) triplinervia nervis secundariis inconspicuis supra anguste insculptis, supra primum furfuracea demum glabrata, subus primum pallide rufo-fulva demum cinerea. Flores in ramulis foliosis plerumque terni sessiles vel vix (ad 1 mm) pedicellati 5-6- (raro 4-) meri. Hypanthium (ad torum) 3.7-4.7 & 3-4 mm.; calyx 1.4-2.8 mm ecoalitus ad ‘ 110 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [voL. 10 apicem vix (ad 0.8 mm) lobulatus, cum dentibus exterioribus 1.d-2.5 & 0.8-1 mm lanceolatis instructus. Petala 12-14 « 5-9 mm, asymmetrice obovata apice obtusa eciliata intus glabra extus modice granuloso-furfuracea. Stamina glabra isomorphiea ; filamenta 3.7-4.8 mm; antherae subulatae curvatae 44.8 mm longae, connectivo sub thecis paulo (ad 0.4 mm) producto postice in calear acutum 0.8—-1 mm longum porrecto. Stylus glaber 8.2-11.3 & 0.5 (basi) —0.2-0.3 (apice) mm; ovarium 3-loculare, dense rufo-granulosum apice in collum integrum 1 mm. longum prolongum ; semina 0.6-1 & 0.8—0.45 mm, laevia pyramidata. | Type: shrub 0.2-0.7 m, petals pink, anthers yellow, occasional in savanna 5 km west of cumbre camp, elev. 1900 m., Cerro de la Neblina, Terr. Amazonas, Venezuela, Jan 6, 1954, Bassett Maguire, John J. Wurdack, & George S. Bunting 37105 (NY). Paratypes (all from the west cumbre of Neblina): Magwire, Wurdack, & Bunting 36927, 37085, 37295, The long exterior calyx-teeth are reminiscent of the genus Meriania; however, the character of the pubescence and the granulose ovaries indicate Graffenrieda. G. fruticosa seems to be the ultimate reduction of a line coming from G. hitchcocku Gleason through G. rufa Wurdack. Graffenrieda reticulata Wurdack, sp. nov. G. fantasticae Schult. & Sm. affinis et in habitu similis, sed foliis trinerviis nervulis bene reticulatis fructibus maioribus. Arbor parva, ramis crassis paulo quadrangularibus (cum cortice laevi sulcato) primum rufo-squamulosis mox glabratis. Petioli 1-1.5 em; lamina glabra 6—10 x 3-6 em, elliptica, apice rotundata vel paulo retusa, basi late acute vel anguste obtusa, trinervia nervis lateralibus maxime 4-8 mm a marginibus distantibus, nervulis supra obscuris subtus leviter elevatis secundariis numerosis tertiaris bene reticulatis. Panicula glabra; flores sessiles cum characteribus generis; alabastra 4-mera, ovario 3-loculari apice rotundato glabro. Fructus urceolatus 53-locularis ca. 8 & 4 mm ad torum ad 2.5—-3 mm econstrictus, calyce vix lobato. Type: tree 5-10 m, fruiting, common in scrub forest between cumbre camp and west escarpment, elev. 1700-1750 m, Cerro de la Neblina, Terr. Amazonas, Venezuela, Jan 13, 1954, Bassett Maguire, John J. Wurdack, & George S. Bunting 37253 (NY). Paratype: bushy tree 3-8 m, buds greenish, occasional in uppermost slope forest above Camp 4, elev. 1600-1700 m, Cerro de la Neblina, Jan 13, 1954, Maguire, Wurdack, & Bunting 37268. G. fantastica has 5- or indistinctly 7-nerved leaves with the tertiary veins not obvious amd an ovoid fruit less than 5 mm long. The paratype collection of G. reticulata had buds sufficiently advanced to establish the genus and flower-mery definitely. Graffenrieda sessilifolia Triana subsp. occidentalis Wurdack, subsp. nov. A subsp. sessilifolia differt petals roseis ovario apice in collum ea. 1 mm longum protracto. Type: shrub or small tree 2-10 m, petals pale pink, anthers pale yellow, oc- casional among boulders on hillside, elev. 1750 m, Cerro Yutaje, Terr. Amazonas, Venezuela, Feb 17-19, 1953, Bassett Maguire & Celia K. Magwmre 35289 (NY). Paratypes (all VENEZUELA, Terr. Amazonas): tree 5 m high, petals pink, anthers yellow, frequent in scrub bush among boulders above Intermediate Camp, elev. 1200 m, Cerro - Yutaje, Feb 4, 1953, Maguire & Maguire 35052; shrub 2 m high, flowers pink, occasional on summit of Cerro Yutaje, elev. 2200 m, Feb 17-19, 1953, Maguire § Maguire 35331; small tree with subcoriaceous shining leaves, petals pink, anthers pale yellow, frequent in subxerophytic 1958] | THE BOTANY OF THE GUAYANA HIGHLAND—PART III 111 low bush on northwest ridge above Camp Yutaje, elev. 1500 m, Cerro Yutaje, Feb 23, 1953, Maguire §& Maguire 35399; tree 2-4 m high, petals pink, occasional on rocks, elev. 1500 m, Cerro Coro-Coro, March 2, 1953, Magwire §& Maguwire 35464; shrub 3 m, flowers pink with yellow centers, Cerro Yavi, elev. 2200 m, March 1-3, 1947, K. D. Phelps g C. B. Hitchcock 55. Typical G. sessilifolia, as represented on Roraima, Ilu-tepui, Ptari-tepui, and Chimanta-tepui, has white petals and ovary apex truncate without collar. On Auyan-tepui exists still another population with pink petals, persistent pubescence on the lower leaf-surface, and truncate ovary apex; this subspecies will be described at a later date. G. sessilifolia is quite variable in leaf size and degree of leaf venation development; the ovary and fruit may be either 2- or 3-celled. Bertolonia venezuelensis Wurdack, sp. nov. Figure 12, b-f. A congeneribus differt antheris basi postice non distincte calcaratis. Caulis repens (raro vix erectus) cum petiolis dense rufo-tomentosus pilis gracilibus flexuosis 1-2 mm longis (ad nodos ad 5 mm longis). Folia in caulibus repentibus alterna jugi uno suppresso (in caulibus raris erectis opposita) ; petiolus plerumque 4-8 em; lamina 6-18 * 3-6 cm, ovata apice per 1l-2 cm caudato-acuminata basi late acuta vel truncata margine integra vel paulo irregulariterque undulata (acumine plerumque_ distincte pinnatilobulato ) breviter 7-plinervia (nervos marginales includens) neryis interioribus 2-10 et 9-17 mm supra basim insertis nervulis supra obscuris subtus leviter distincteque elevatis, supra modice strigulosa pilis gracillimis 5-8 mm? ca. 1 mm longis pallidis, subtus in superficie glabra in nervis primariis densissime rufo-strigosa (pilis ut in caulibus) in secundariis modice strigulosa tertiariis glabris sed cum glandulis pallidis clavatis armatis. Pedunculus communis erectus (ad ramos 10-15 cm) primum modice rufo-strigulosus demum subglabratus, ca. medium cum nodo; rami plerumque umbellati ad anthesim 2-3 cm longi secundiflori ad nodos bibracteolati, bracteola sessili submillimetrali basi cum glandulis duobus nigris globosis sessilibus subtenta. Pedicelli 2-4 mm cum hypanthio sparse strigu- losi pilis gracilibus ca. 0.1-0.2 mm longis. Flores 5-meri. Hypanthium 1.7 (ad torum) & 2.5 mm; calyx 0.4 mm coalitus, lobis 0.8 & 2.2 mm, basim versus paulo imbrieatis late ovatis apice truncatis extus tuberculato-dentatis dente ca. -1 mm longo non vel vix projecto. Petala 9.5-10 < 7.5-8.5 mm obovata vel subrotundata paulo asymmetrica apice rotundata glabra eciliata. Stamina isomorphica; filamenta 3.2-3.7 mm longa; antherae 2.7-3.2 mm longae paulo subulatae, connectivo exappendiculato sub loculis imperspicue (0.1-0.15 mm.) producto. Stylus 5-5.5 & 0.2-0.8 mm.; stigma punctiforme; ovarium triloculare triquetrum glabrum apice in lobulos tres obtusos carnosos 0.6 mm altos extensum. Type: prostrate herb, petals very pale pink inside, darker without, stamens yellow, locally frequent on escarpment slopes east of Camp 3, elev. 1600 m, Cerro de la Neblina, Terr. Amazonas, Venezuela, Jan 24, 1954, Bassett Magwre, John J. Wurdack, & George S. Bunting 37370 (NY). Paratypes (same locality) : Dee 27, 1953, Maguire, Wurdack, & Bunting 36951. 36947. B. venezuelensis represents the first record for the genus outside of south- eastern Brazil. In habit, the Neblina species agrees quite well with several Brazilian species, but the combination of densely rufous-pilose stems and pubescent acuminate leaves are sufficient vegetative distinctions. The connective being not at all prolonged into a dorsal calear (albeit prolonged slightly below the thecae to the filament insertion) seems unique. ‘ [vou. 10 12 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN , “s (i wai HIN SS Fi Fa 4 = Fic. 12. a, Macrocentrum gracile, habit, X 0.5: x 0.5; ¢, flower, X 4; d, stamen, X 2; e, pistil, X 2; f, ovary top view, X 2. cP re X SS [[Evare », YQ & b-f, Bertolonia venezuclensis. b, habit, 1958 | THE BOTANY OF THE GUAYANA HIGHLAND—PART III 113 Macrocentrum gracile Wurdack, sp. nov. Figure 12, e. M. fruticoso Gleason affinis sed foliis maioribus et tenuioribus basi distincte cordatis pedicellis longioribus petalis minoribus connectivo antherarum dorsali minore. Plantae herbaceae suberectae, caulibus ad 30 cm longis, quadratis vel sub-4- alatis glabris. Folis opposita in quoque jugo aequalia vel subaequalia; petiolus 2.8 em, glaber; lamina tenuissima 4-12 * 3-7 em, ovata vel oblongo-ovata, apice breviter (ca. 1 cm) hebetiacuminata, basi breviter (ad 1 cm) cordata lobis basilaribus plerumque imbricatis, 5—7-nervia nervis primariis gracilibus secundariis indistinctis, ad marginem integra vel vix denticulata ciliata ciliis patentibus ca. 1-2 mm longis et 1-2 mm inter se distantibus, supra sparse strigosa pilis gracilibus 1-2 mm longis 0.5-3 mm?, subtus in superficie venisque primarus elabra in venulis (praecipue marginem versus) sparsissime — strigulosa. Inflorescentia laxa glabra pedunculo gracili 6-8 em longo ramis_ primariis umbellatis plerumque 2-3 usque 4 em longis, ramulis umbellatis 3 vel 4 in quoque ramo 1-3 em longis 2—4-floris. Flores secundi 4-meri glabri; pedicelli graciles, d-10 mm. longi. Hypanthium 3 X 1.2 mm; ealyx vix (0.1-0.2 mm) coalitus, lobis 0.4 & 0.9 mm late triangularibus obtusis extus cum dente 0.4 mm longo vix (0.2 mm) projecto armatis. Petala 3.6 & 1.4 mm anguste obovata acuta. Stamina isomorphica; filamenta 1.8 mm longa; antherae loculi 2.5 mm longi, apice attenuati, connectivo sub loculis ad insertionem filamenti non producto sub filamento dorsaliter in caudam 0.4 & 0.1 mm hebeti-acutam producto. Ovarium biloculare ; stylus 4.5 & 0.1 mm; stigma punctiforme; semina 0.28 & 0.2 mm, ovoidea unilateraliter truncata minute tuberculata. Type: delicate annual with white flowers, infrequent under overhanging rocks on summit of southeast-facing escarpment, elev. 700 m, Mt. Ebeni, British Guiana, Oct 15, 1951, Bassett Maguire 32117 (NY). M. fruticosum has smaller thicker leaf-blades with usually truncate bases, pedicels only 1-3 mm long, petals 7-7.5 mm long, and larger anthers with relatively longer dorsal appendage. Macrocentrum stipulaceum Wurdack, sp. nov. M. parvulo Gleason affinis sed foliis plusminusve acuminatis distincte denticulatis, ramulis ad nodos cum processibus reniformibus stipulaceis armatis. Herba parva ad 15 em alta, caulibus glabris. Folia opposita aequalia vel paulo inaequalia; petiolus gracilis glaber 5-15 mm, cum processu reniformi stipuliformi sparse glanduloso-strigoso 1-2 * 1.3-2.5 mm subtentus processibus subpersistentibus 2 (—3?) per nodum; lamina tenuissima 15-40 x 10-20 mm, ovata apice gradatim hebeti-acuminata basi paulo (1-3 mm) cordata, 3—5-nervia, ad margines obtuse denticulata dentibus ca. 0.1 mm altis 1-2 mm inter se distantibus in dentis sinu ciliata ciliis gracilibus 0.5-1 mm longis, supra et subus sparse strigulosa setis gracilibus ca. 1 mm longis ca. 1/mm? praecipue praeter venas venulasque. Pedunculi terminales 1-3 em longi, glabri, 1- vel plerumque 2-fiori. Flores ignoti; pedicelli fructuum ca. 5 mm lonei. Hypanthium (ad torum) ca. 4.5 mm longum paulo 8-venosum; calycis lobi interiores quattuor obtusi, ca. 0.7 1.3 mm, dente exteriore acuto ca. 0.4 mm projecto, setis paucissimis elanduloso-capitatis ad 1 mm longis extus in calycis lobis notatis. Semina cum aspectu M. gracilis, ca. 0.8 & 0.6 mm, minute tuberculata. Type: delicate annual, locally frequent in dry places under overhanging rocks on montane slopes of Mt. Ayanganna., elev. 1000-1500 m. British Guiana, s ; ‘ 114 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [voL. 10 Feb 5-6, 1955, Bassett Magwre, W. M. C. Bagshaw, & Celia K. Maguire 40582 (NY). M. parvulum has shorter stems, obtuse to subacute leaf-blades with obtuse to broadly acute bases, and no stipuliform cauline appendages. The leaf shape, lack of stipules, and many-flowered inflorescences will distinguish M. montanum Gleason and depauperate specimens of M. cristatum Tr. and M. fruticosum Gleason. Macrocentrum longidens (Gleason) Wurdack, comb. nov. Diolena longidens Gleason, Mem. N. Y. Bot. Gard. 8: 140. 1953. Macrocentrum repens (Gleason) Wurdack, comb. nov. Diolena repens Gleason, Bull. Torrey Club 75: 541. 1948. For both these taxa I have now been able to examine buds sufficiently mature to show the stamen structure. The affinity of both with the next-described species is evident, in aspect as well as in floral details; all three have a ciliate short ovarial collar, the same type of anther and stamen appendages, and the same kind of hypanthial pubescence. These transfers eliminate the genus Diolena from the Guayana Highland flora. MW. repens is now also known from the slopes of Ptari-tepui (Steyermark 59461, Maguire & Wurdack 33928) and from Mt. Avanganna (Maguwire, Bagshaw, & Magwire 40584-A). Macrocentrum neblinae Wurdack, sp. nov. M. longidenti (Gleason) Wurdack valde affinis sed plerumque robustius, foliis maioribus in proportione angustioribus apice anguste acutis sepalorum lobis interioribus ovatis hebeti-acutis. Herba basi lgnosa alt. ad 60 em superne ramosa, ramis teretibus glabris. Folia opposita in quoque jugo valde dimorphica primum cum glandulis pallido- rubris sparse ornata demum marginibus exceptis glabra; folia minora sessilia ovate acuta ad 8 & 3 mm uninervia margine sparse denticulato-ciliata; folia maiora brevipetiolata (petiolo plerumque ca. 1 mm longo) 2-4 0.5-1.3 em, lanceolata apice acute basi cuneata paulo (0.5-1 mm) triplinervia, marginibus de medio apicem versus sparse denticulatis, denticulis 1-2 mm inter se distantibus uniciliatis cilio appresso. Flores 5-meri solitarii, pedicello ca. 2-8 mm longo, dense elanduloso-puberulo basi bibracteolato bracteolis lanceolatis g@labris. Hypanthium (ad torum) 2-2.8 & 2.2-2.7 mm modice glanduloso-puberulum pilis ca. 0,5 mm longis. Calyx vix (0.2 mm) coalitus. lobis interioribus 2-2.0 1.6-1.9 mm. ovatis hebeti-acutis sparse elanduloso-puberulis margine glanduloso- ciholatis, dentibus exterioribus aristatis arista 2-2.5 mm eminenti (in fruectu ad 3 mm) basim versus parce glanduloso-scabridula. Petala (paulo immatura) 4.7 & 2.7 mm ovata acuta breviapiculata elabra. Stamina inconspicue dimor- phica; filamenta matura desunt; antherae 2.6 mm longae, basi 0.5 mm latae, paulo subulatae poro 0.2 mm lato, connectivo ad insertionem filamenti 0.3 mm producto, appendice dorsali 0.6 mm longa hebetiacuta. Stylus 4.5 & 0.2-0.3 mm; stigma paulo capitulatum 0.35 mm latum; ovarium 3-loculare glabrum apice 1m collum breviciliatum 0.5 mm productum; semina 0.5 < 0.3 mm, laevia pyrami- data paulo aneulata. Type: flowers white, locally common on boulders in stream courses in talus forest between Camps 2 and 3, elev. 250-650 m, Cerra de la Neblina, Terr. Ama- zonas, Venezuela, Dee 22, 1953, Basset Maguire, John J. Wurdack, & George SW. 1958 | THE BOTANY OF THE GUAYANA HIGHLAND—PART III 115 Bunting 36802 (NY). Paratype: occasional on and around rocks in forest south- east of Camp 3, elev. 900 m, Cerro de la Neblina, Jan 24, 1954, Maguire, Wur- dack, & Bunting 37364. The leaf-blades of M. longidens are only 12-17 & 7-9 mm and elliptic or scarcely ovate-elliptic with broadly acute apices; the interior calyx lobes are semi-circular. Macrocentrum maguirei Wurdack, sp. nov. M. anychioidi Gleason affinis sed foliis longioribus oblanceolatis, petalis an- therisque minoribus, staminum appendice dorsali longiore. Herba exigua ad 15 em alta superne ramosa caulibus cum internodiis glabris subquadrangulatis nodis setosis setis robustis ca. 1 mm longis. Folia isomor- phica; petiolus 1-3 mm longus; lamina 7-15 X 1.5-4 mm, oblanceolata apice anguste rotundata basi attenuata uninervia sparse (praecipue apicem versus) ciliata ciliis ad 1.5 mm longis et 1-3 mm inter se distantibus, supra et subtus primum sparse rubro-glandulosa supra demum glabrata. Flores solitarii 4-meri; pedicellus sub bracteolis ca. 5 mm longus, supra 0.5—-0.7 mm; bracteolae 0.9 0.2 mm. Hypanthium 1-1.3 x 1.7-2 mm, sparse rubro-glandulosum breviter 8- alatum alis apice unisetosis seto ca. 1 mm longo. Calyx 0.3 mm coalitus, lobis 1.8 X< 1 mm late semicircularibus paulo emarginatis. Petala 5.5 & 3 mm, oblongo-obovata apice subtruneata glabra. Stamina isomorphica; filamenta 2.8 mm; antherae thecae oblongae 1.1 & 0.3 mm, poro 0.1 mm diam., connectivo sub loculis (ad filamenti insertionem) inconspicue (0.1 mm) producto, appendice . dorsali 0.6 mm longa hebetiacuta. Stylus 2.9 * 0.3-0.5 mm; stigma 0.4 mm latum,*vix capitulatum; ovarium 3-loculare, apise truncatum; semina 0.5 0.3 mm, late pyramidata minute muriculata. Type: flowers pink, infrequent in moist shaded places at base of waterfalls at head of Cand Yutaje, elev. 1900 m, Cerro Yutaje, Terr. Amazonas, Venezuela, Feb 17-19, 1953, Bassett Maguire & Celia K. Maguire 35298 (NY). M. anychioides has a leaf-blade leneth-width ratio of 2-2.3 (rather than 34.5), much larger petals, larger linear anthers, and dorsal connective ap- pendage only 0.2-0.3 mm long. Salpinga paleacea Wurdack, sp. noy. Figure 13. A congeneribus differt petiolis venis foliorum subtus cum pilis paleaceis grandibus armatis. Subherbacea simplicicaulis, caule subprostrato robusto ad 20 em longo, ea. 1 cm crasso, ad nodos densissime paleaceo-setoso (setis ca. 10 mm longis, basi 1-1.5 mm latis) alioqui glabro. Petiolus 6-15 x 0.2-0.4 em, modice paleaceo- setosus et primum gracili-strigosus pilis gracilibus mox caducis; lamina 12-20 < 9-14 em, late elliptica vel suborbicularis, apice rotundata vel brevissime abrup- teque obtuso-apiculata, basi paulo cordata, 7-nervia nervis et nervulis supra paulo insculptis subtus elevatis nervis secundariis 5-10 mm inter se distantibus, margine dense setuloso-ciliata setulis subincurvis 2-3 mm longis, supra primum dense villosula mox glabrata, subtus inter nervulos glabra ad nervos primarios et secundarios modice appresso-setosa (setis longo-attenuatis ad 10 * 0.2 mm) et ad nervos nervulosque gracili-strigulosa pilis laxis paulo caducis. Pedunculus 20-30 em longus, ad nodum subbasilarem dense paleaceo-setosus alioqui primum gracili-strigulosus demum glabratus. Flores 5-meri, in ramis umbellatis ad pedunculum summum secundi; rami divaricati basi modice paleaceo-setosi pri- paces PIE PES. [vou. 10 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN \ Fae S Eee BS ee ra eee “ie A eo 6. Pe LAR Mem = WG PX kA Vie Ae D Fig. 13. _Salpinga paleacea. mens, X 4. a, habit, X 0.5; b, flower, X 2; e-d, large and small sta- £9584. > THE BOTANY OF THE GUAYANA HIGHLAND—PART III ha mum cum pilis gracilibus crispulis modice armati (pilis demum paulo caducis) ad anthesim 1—2 em longi demum ad 3 em longi. Pedicelli 1-2 mm longi; hy- panthium (ad torum) 3.3 x 2.5 mm, sparse gracili-strigulosum; calyx supra torum 1.3 mm coalitus, lobis 0.5 & 2.9 mm, rotundatis extus cum dente 0.2 mm alto non eminenti armatis. Petala ecilata glabra, 10.8-11.8 & 7-8 mm, asym- metrice obovata apice rotundata. Stamina dimorphica. Staminum maiorum: filamentum 4.4 mm longum; thecae descendentes subulatae 5.2 mm longae, con- nectivo postice basi brevissime (ad 0.15 mm) calearato et in appendicem an- therae parallelem 2 & 0.4-0.8 mm producto appendice apice breviter (0.4 mm) bilobata lobis rotundatis. Staminum minorum: filamentum 4 mm longum; thecae descendentes subulata 3.2 mm longae, connectivo postice indistincte calcarato et in appendicem antherae parallelem 1.5 & 0.5 mm producto appendice apice rotundata non lobata. Stylus 6.8 & 0.8— (apicem versus) 0.2 mm, glaber; ovari- um glabrum, 3-loculare, apice truncatum; hypanthium fructiferum ca. 3 mm longum, 7-10 mm pedicellatum teres (indistincte 10-striolatum), calyce divari- cato 6-7 mm lato; semina 1 X* 0.4 mm laevia pyramidata. Type: perennial herb, frequent in deep humus at base of rocks, summit of upper east-facing escarpment of Mt. Ebeni, elev. 700 m, British Guiana, Oct 15, 1951, Bassett Maguire 32119 (NY). None of the other known species of Salpinga has a paleaceous indument. In fruit shape, S. paleacea is intermediate between the southeast Brazilian species (with trigonous fruit) and the remainder of the genus (with long hypanthia). The stamens resemble those of S. secunda Schr. & Mart. ex DC. and S. maguwirei Gleason (and probably S. ciliata Pilger) in having a very minute basal spur on the connective rather than the prominent spur pictured by Triana (Trans. Linn. Soc. 28: pl. 6, f. 80. 1871) for 8S. margaritacea (Naud.) Tr. and S. longifolia (Cham.) Tr. The habit of S. paleacea is rather reminiscent of that of Tococa platyphylla Benth. APOCYNACEAE” Ambelania oleaefolia Monachino, sp. noy. Figure 14. Frutex glaber; foliis oppositis coriaceis; petiolis ca. 5-7 mm longis; laminis ellipticis, 3-10 em longis, 0.7-3 em latis, non nitidis, ad basin obtusis, ad apicem rotundatis vel obtusis vel raro acutis; nervis secundariis utroque ca. 25, supra debilibus, subtus plerumque obscuris; inflorescentiis terminalibus 3—6-floris ; sep- alis ovatis obtusis 1.5-2.7 mm longis, glabris, squamellis numerosis; corolla mag- na, extus glabra, tubo ca. 2 em longo, intus pubescenti, lobis oblongo-oblanceo- latis ca. 2-3 em longis, 1 em latis; staminibus supra basin tubi corollae 5 mm insertis ; antheris sessilibus ca. 4.8 mm longis paene usque ad basin dehiscentibus, lobis basalibus rotundatis non patentibus; ovario glabro; fructibus ELE OLReS usque ad 4 em longis, 2—2.5 em latis; seminibus numerosis. Shrub 0.2-2 m tall, glabrous, the stem subterete; leaves opposite, rigidly coriaceous; petiole about 5-7 mm long; blade elliptic, sometimes narrow, obtuse at the base, rounded to obtuse or rarely acute at the apex, 3-10 em long, 0.7—3 em wide, rather dull, slightly paler beneath, the lateral nerves mostly obscure 22 By Joseph Monachino. ‘ 118 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [voL. 10 7 [CLARE | Fic. 14. a-d, Ambelania oleaefolia. a (Maguire. g& Wurdack 34512), stem with leaves and flowers, X %; b (Maguire, Cowan & Wurdack 30553), stamens, X 3; ¢, (Maguire, Cowan & Wurdack 30474), fruit, *X 4; d (same coll.), seed, K 2%. e-h, Lacmellea pygmaea (Maguire § Wurdack 34507). e, stem with leaves and flowers, X 2; f, fruit, K 2%; g, stamens, X 5; - h, style and stigma, * 10. 1958 | THE BOTANY OF THE GUAYANA HIGHLAND—PART III 119 beneath, faint on the upper side, the principal secondaries about 25 pairs with an average distance of 2-4 mm apart near the middle of the blade; inflorescence a terminal, mostly 3—6-flowered cyme, the common peduncle up to 1.5 em long, the pedicels up to 8 mm long, with deltoid or ovate small bracts atthe base; sepals persistent, broadly ovate, obtuse at the apex, 1.5-2.7 mm long, glabrous, non-ciliate, intracalycine squamellae numerous, ovate or oblong, about 0.4—0.5 mm long; corolla white, fragrant, hypocrateriform, large, glabrous outside, the corolla-tube straight or slightly curved, about 2 em long, glabrous at the base within but densely pubescent above the insertion of the stamens up to the corolla-lobes, the hairs cylindrical becoming clavate toward the orifice of the tube, the corolla-lobes sinistrorsely convolute, oblong-oblanceolate, about 2—3 em long, 1 em wide, glabrous; stamens inserted about 5 mm from the base of the corolla- tube, the anthers sessile, lanceolate, about 4.8 mm long, dehiscent nearly to the base, the terminal sterile membranous acumen about 0.3 mm long, the basal lobes rounded, not spreading; ovary synearpous, conical above, somewhat rounded at the apex, about 1.7 mm long, glabrous, the style (of a mature flower) 2.3 mm long to the incrassate ring, the crested capitulum about 0.5 mm long, the annular disk closely cohering to the base of the ovary, thin, low, irregular, faintly undu- late, less than 0.5 mm high; fruit a berry, ellipsoid, sometimes narrowed toward the base, rounded, up to 4 em long, 2—2.5 cm in diameter, smooth or verrucose ; seeds numerous, irregularly oblong-elliptic, 5-7 mm long, 3-4 mm wide, spongy, minutely roughened with protuberances and microscopically pitted; embryo linear, surrounded by a thin fleshy endosperm ; cotyledons oval, about 1 mm long; radicle about 4 mm long. Type: Bassett Maguire & John J. Wurdack 34512, Alto Rio Orinoco, Terri- torio Amazonas, Venezuela, frequent at edge of Savanna No. 3, northwest base of Cerro Yapacana, elev. 150 m, March 17, 1953, shrub 0.2-0.8 m, flowers white, fragrant (holotype NY). Paratypes: VENEZUELA: Amazonas: Maguire, Cowan §& Wurdack 30474, Cerro Yapa- cana, Rio Orinoco, occasional, savanna at base of Cerro Yapacana, 175 m alt., Dee 31, 1950, sprawling shrub to 5 dm, flowers white, fragrant; unicate. Maguire, Cowan §& Wurdack 30553, Cerro Yapacana, Rio Orinoco, occasional, Savanna No. 3, Jan 1, 1951, shrub to 2 m, flowers white; unicate. Maguire & Wurdack 34504, Alto Rio Orinoco, occasional in Savanna No. 2, northwest base of Cerro Yapacana, elev. 150 m, March 17, 1953, shrub 0.2—0.8 m, flowers white, fragrant. This species is readily separated from the dozen previously known species of Ambelana. The long corolla-tube, pilose within, would suggest comparison with A. macrophylla Muell.-Arg. and A. grandiflora Huber in the revision of the venus by Monachino (Lloydia 8: 109-130. 1945). The leaves in our newly dis- covered plant are much smaller. The leaf size is closer to that of A. lava ( Benth.) Mnuell.-Are., but here the resemblance even in foliage stops at size alone. It is interesting to note that A. oleaefolia grows in savanna country, an habitat quite different from that of A. Jara. It does not have the stem swollen at the base like the riparian A. lara. Aspidosperma album (Vahl) R. Ben. ex M. Pichon? Maguire §& Wurdack 35663, Rio Guainia, Amazonas, along Cano Pimichin below Pimichin, elev. 125-135 m, April 14, 1953, slender tree 10 m, fruit brown. Only a fruiting specimen is available. The matured follicle is oblong-obovate and much smaller than has been described for either this species or A. spruce- anum. Benth. ex Muell-Arg., about half the size, 4 em long, 2 cm wide. The stipe ‘ 120 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 is a little over 1 em long. The seeds are numerous, oblong to triangular, nar- rowly papery-winged, 2-2.5 em long, 1.3-1.7 em wide; cotyledons orbicular, about 1.38 em wide, the sinus closed, the radicle 3 mm long, not surpassing the lobes of the cotyledons. The plant is named A. album rather than A. spruceanum because the petiole is longer than that of the latter, 1.5-2.5 cm long, and the venation on the underside is slightly more expressed. Woodson (Ann. Missouri Bot. Gard. 38: 196. 1951) hypothesizes possible hybridization to account for variability in A. album. Aspidosperma decussatum Woods. The fruit is now known (7. Lasser 1805, Kavanayén, Bolivar, Mr 1946). Follicle stipitate, asymmetrically oval, narrowed at the base, about 7 em long, 4.5 em wide, velvety-pubescent, the ridge about 2 em from the nearest margin, the stipe 1.5 em lone; seeds many, orbicular, papery-winged, about 4 em wide; cotyledons orbicular, about 1.5 em wide, the sinus closed; radicle 3 mm long, not surpassing the lobes of the cotyledons. This opposite-leaved species has been collected only in the area of the type locality, Ptari-tepui, Kavanayén, and Chimanta. Aspidosperma exalatum Monachino, sp. nov. Figure 15. Arbor; ramulis primo puberulis, denique glabris; foliis alternis coriaceis olabris; petiols 1—1.5 em longis; laminis oblongis, (6—) 10-13 em longis, (3—) 4-6.5 em latis, ad basin obtusis vel rotundatis, ad apicem rotundatis emarei- natisque, subtus pallidis; nervis secundarus utrinque prominentibus, reticulo venulorum prominulo; fructibus sessilibus crasse lignosis inaequaliter ovatis, \-10 em longis, 3.5-4.5 em latis, tomentulosis minute rugulosis, ad apicem apicu- latis; seminibus suboblongis vel ovatis, 4-5.5 em longis, 2.5—3 em latis, non membranaceo-alatis, marginibus crassiusculis angustisque; cotyledonis subob- longis, ca. 4 em longis, 2 cm latis; radicula ca. 5 mm longa, lobos cotyledonorum ‘non excedenta. Tree 5-7 m tall, the branchlets moderately stout, puberulent when young, becoming glabrous; leaves alternate or approximate, rigidly coriaceous, complete- ly glabrous (old leaves) ; petiole dark, 1-1.5 em long; blade oblong, obtuse or rounded at the base, rounded and emareginate at the apex, thick-rimmed at the margins, (6—) 10-13 em long, (38—) 46.5 em wide, paler beneath than above but not markedly elaucous, the secondary nerves raised on both surfaces, the princi- pal ones averagine 5-7 mm apart near the middle of the blade, the reticulation of veinlets prominulous; flowers lacking; sepals (in fruit) ovate-lanceolate, white-tomentulose outside, about 2 mm lone; follicle thickly woody, asymmetri- cally oval, sessile, apiculate at the apex, 5-10 em long, 3.5-4.5 em wide, closely brownish or rusty-tomentulose, minutely roughened, the ridge about 2 em from the nearest margin; seeds few in each follicle, smoke-colored, centrally tinted with bluish-green, oblongish varying to ovate, not papery-winged, the margins rela- tively thick and narrow, less than 1 cm beyond the embryo; cotyledons oblonge- ish, cordate or emarginate at the base with a close sinus, about 4 em long, 2 em wide; radicle about 5 mm long, not surpassing the lobes of the cotyledons. Type: Bassett Maguire & John J. Wurdack 34611, Alto Rio Orinoco, Terri- torio Amazonas, Venezuela, along Yapacana caho between laguna and Rio Ori- noco, elev. 150 m, March 20. 1953, tree (holotype NY). Paratype: VENEZUE- 1958 | THE BOTANY OF THE GUAYANA HIGHLAND—PART III 121 LA: Amazonas: Magwre & Wurdack 34909, Rio Casiquiare and tributaries, occasional along Rio Pacimoni between Canos Arapacua and Boridahari, elev. 110 m, April 7, 1953, spare tree 5-7 m, fruit brown. The leaves of A. exalatum were not matched with any species heretofore de- scribed. The venation somewhat recalls that of A. verruculosum Muell.-Arg., but the secondaries are much further apart. The reduced thickened wing of the seed is a feature which was not noted for any species of Aspidosperma by Woodson in his monograph of the genus. However, A. Ducke (Anais Acad. Bras. Ci. 27: 382. 1955) mentions such a wing for A. nitidwm Benth. ex Muell.-Arg. Fruiting material is not available for many species and consequently it remains to be seen how rare exalate seeds are in the genus. Aspidosperma glaucum Monachino, sp. nov. Figure 15. Arbor; ramulis glabrescentibus; foliis alternis coriaceis subglabris; petiolis ca. 2 em longis; laminis oblongo-obovatis vel obovatis, 10-12 em longis, (3—) 4-6 cm latis, ad basin cuneatis, ad apicem obtusis, supra nitidis, subtus glaucis; nervis secundaris debilibus; venis obscuris; fructibus crasse lignosis ovali-pyri- formibus, ca. 12 cm longis, 8.5 em latis, glabris minute rugulosis; seminibus orbicularibus, 7-8 em latis, membranaceo-alatis ; cotyledonis late ovatis, ca. 3 em longis, 2.5 em latis; radicula ca. 5 mm longa, lobos cotyledonorum excedenti. Tree 20 m tall, essentially glabrous, the branchlets moderately thick, brown- ish, glabrescent, the buds grey-tawny-puberulent; leaves alternate or approxi- . mate; petiole 0.8—2.3 em long, brownish, sometimes sparsely appressed-puberulent particularly toward the base; blade coriaceous, oblong-obovate or obovate, cune- ate at the base, very short-acuminate or rather apiculate at the apex, revolute at the margins, 10-12 em long, (3—-) 4-6 em wide, essentially glabrous, a few scattered hairs (up te about 0.17 mm long) sometimes present particularly along the midrib, shining above, strikingly glaucous-colored beneath (covered with scurfy material that can be scraped off), the secondary nerves faint, not raised, the principal ones averaging 5-10 mm apart near the middle of the blade, the veins obscure; flowers lacking; follicle (unattached) thickly woody, oval- pyriform, about 12 em long, 8.5 em wide, glabrous, the surface minutely rough- ened, the ridge 3.5 em from the nearest margin; seeds papery-winged, more or less orbicular, 7-8 em wide; cotyledons broadly ovate, shallowly cordate at the base, about 3 em long, 2.5 em wide; radicle 5 mm long, surpassing the lobes of the cotyledons. Type: Bassett Magwre & Louis Politi 28255, Cerro Sipapo, Territorio Ama- zonas, Venezuela, cumbre, Savanna Camp to Cano Profundo and Kast Terrace via Cano Negro, elev. 1600 m, Jan 8, 1949, tree 20 m high, 20 em diam., leaves elaucous beneath, seed broadly membranous winged (holotype NY). In its coriaceous, glabrous, glaucous-colored leaves, A. glawcum recalls A. excelsum Benth. (altogether different in its spinose-verrucose follicles), the pallor on the under side of the leaves of the former being more marked. The leaves of our new species seem to differ further in being apiculate at the apex, in having revolute margins and secondary nerves appreciably fainter. The buds are tawny- tomentulose, not grey as in A. excelsum. As indicated above, A. glaucum is easily and with certainty separated from A. excelsum by fruit character. When, however, it is compared with A. schultesu Woodson, similar in its heavily coriaceous leaves with whitened under side and ‘ 122 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [voL. 10 ’ J fli BARDS x v4) iI NAS ‘ | HAN : es \ \) \ \\ \\\ ‘ x CLM, yi NTN\ PH YZ : \\\ a SA Ss: Wy N Se N = Re Ne NG N z moe He Fig. 15. a,b, Aspidosperma exalatum (Maguire & Wurdack 34611): a, leaf and fruit, x %; b, seed, X 1. e-e, Aspidosperma glaucum (Maguire & Politi 28255). ¢, fruit, K %; d, seed, KX %; e, leaf, K %. f,g, Aspidosperma neblinae (Maguire, Wurdack § Bunting 37084). f, leaf, KX %; g, flower, X 5. h-—j, Paralyxia macrophylla (Guppy 7494). h, fruit, * WY; 1, seed, X< 46% .j, embryo, lL. . 1958 | THE BOTANY OF THE GUAYANA HIGHLAND—PART III 123 immersed venation, there is difficulty. The two species are known only from the types. The material is essentially incomparable, as one is in fruit, the other in flower. The stems of the branchlets are blackish in A. schultesiwi, the petioles much longer, 4-4.5 em long, black, the blades relatively longer and narrower, not as hoar-frost-white beneath but rather the color of those of A. excelsum. The under side of the leaves is covered for the greater part with very short grey appressed hairs, while those of A. glawcum are either glabrous or have scattered longer hairs along or near the midrib. The young leaves and calyx of A. schultesw have a sulphur-colored indumentum; and whereas the buds of A. glau- cum are also colored, they are of a different hue, a shade of pale tawny, and the indumentum consists of longer hairs. From consideration of the different habi- tats of the two plants, it would seem unlikely that they represent variations of the same species. A. schultesii was collected on a low ridge separating two watersheds, headwaters of Ira-Igarape and Igarape Abiu, affluent of Rio Taraira, Amazonas, Brazil, in a typical balata-type forest, in a region of endemism (com- munication from Dr. Schultes). A leaf of the type was sent to Heber W. Youneken for microscopic examina- tion. Dr. Youngken wrote: ‘‘I have made a microscopic examination of portions of the leaf and find that the scurfy material which can be scraped off from the lower surface consists largely of outgrowths of the epidermal cells as conical to branched, unicellular hairs, the branched type occurring in several forms, the main stalk frequently bifurcate at its distal end into straight or curved sharp- pointed branches, another form resembling somewhat the antlers of the deer or ‘antelope. Many of the hairs possess an indistinct lumen; in others it is ex- tremely narrow, and the walls are greatly cuticularized. Some cuticular scales also oceur.”’ Aspidosperma neblinae Monachino, sp. noy. Figure 15. Arbor; foliis alternis primo subtus tomentulosis dein puberulis; petiolis ca. 1 cm longis; laminis obovatis, 7-11 em longis, 3.5—5 em latis, ad basin cuneatis, ad apicem rotundatis, venis prominulis; floribus congestis; pedicellis quam calyce brevioribus ; sepalis 3-4 mm longis, extus tomentulosis; corolla glabra, tubo angu- lato, 4.3 mm longo, lobis lineari-lanceolatis acutis, 3.7 mm longis; staminibus media tubi corollae insertis; antheris 0.95 mm longis; ovario glabro; fructibus ovali-pyriformibus velutinis, 7-9 em longis, 3.5—6 em latis, stipite 1.5—2 cm longo ; seminibus plusminusve orbicularibus membranaceo-alatis, usque ad 6 cm latis; cotvledonis orbicularibus, ca. 2 em diam. Tree 5-15 m tall, the stems of the branchlets moderately stout, closely to- mentulose when young; leaves alternate (rarely subopposite), coriaceous, closely orey-(faintly tawny-) tomentulose beneath when young, becoming merely puberu- lent; petiole short, about 1 (-1.5) em lone, dark-colored ; blade oblong-obovate or obovate-cuneate at the base, rounded to subemarginate at the apex, (6-) 7-11 cm long, 3.5-5 em wide, the venation somewhat prominulose, the midrib little raised on the upper surface; inflorescence corymbiform; flowers white, crowded, the pedicel shorter than the calyx; sepals acutish, 3-4 mm long, closely grey-tomentu- lose outside, hairy within on the upper part; corolla glabrous, the tube angulate, slightly exserted from the calyx, 4.3 mm lone, the lobes spiral-contorted in aesti- vation, sharply acute, linear-lanceolate, about as long as the corolla-tube, 3.7 mm long; stamens inserted at about the middle of the corolla-tube, the anthers 0.95 mm long; ovary glabrous; fruit stipitate, woody, oval-pyriform, flattened, grey- % 124 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [voL. 10 tawny-velvety-tomentulose, the stipe 1.5—-2 em long, the body 7-9 em long, 3.5—-6 cm broad, the ridge 1.5-2 em from the nearest margin; seeds papery-winged, more or less orbicular, varying in size, 2.5-6 cm in diameter; cotyledons orbicu- lar, about 2 cm broad, cordate at the base with closed sinus; radicle 4 mm long, not surpassing the cotyledon-lobes. Type: Bassett Magwre, John J. Wurdack & George S. Bunting 37084, Cerro de la Neblina, Rio Yatua, Teritorio Amazonas, Venezuela, in scrub forest alone escarpment west of Cumbre Camp, elev. 1700-1800 m, Jan 6, 1954, bushy tree dm, flowers white (holotype NY). Paratypes: VENEZUELA: Terr. Amazonas: Maguire, Wurdack g¢ Bunting 37245, Cerro de la Neblina, Rio Yatua, occasional along west escarpment edge 2 km north of Cumbre Camp, elev. 1800 m, Jan 12, 1954, tree 8 m,-fruiting; unicate. Maguire, Wurdack, & Bunting 37284, Cerro de la Neblina, Rio Yatua, frequent in upper slope forest above Camp 4, elev. 1500-1700 m, Jan 13, 1954, tree 6-15 m, fruit green. A. neblinae belongs in the series Nobiles Woodson and suggests alliance with A. steyermarki Woodson, and less so with A. decussatum Woodson. Aspidosperma sp. aff. A. neblinae Monachino. Cowan §& Wurdack 31166, Serrania Part, Rio Part, Cano Asisa,, Rio Ventuari, Amazonas, Venezuela, occasional, cumbre just south of head of valley of Camp Cano, sabanita and scrub forest, 2000 m alt., Feb 2, 1951, tree 4-10 m tall, fruit green, abundant white latex. Material with flowers past anthesis and matured fruits is available. As far as can be seen in the absence of flowers, the plant differs from . Be igo) Con Wekc \Y a? Ss , \) Bree : ty S) Message te leases ) ivy cy > S co \ ecaee ale rose) y Wee Ogres fon SS Se SISA Nea He (\ 1} \ 280 Pe 1sSa T } WANA DSSS awa S Case: ee eee o ; lS: y Disk PRN) SAIC fo) ans EAN Xe sennene: a Ope, YS 0) 0 OS0%c7 O85 O° 620 O 160 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [voL. 10 ‘ has a ‘‘megaphytic’’ habit of growth; that is, a single thick unbranched stem bearing a rosette of leaves is formed. In the stem with secondary growth shown in figure 3, the high degree of cork development is shown, successive periderms being indicated. This condition is in contrast to the relatively modest develop- ment of secondary xylem. Sclereid nests are relatively infrequent in the cortex, and become included in the cork during periderm formation. Bundle caps occur, although intervening parenchyma never becomes sclerified to connect them as it does in Stenopadus. In the secondary phloem files or patches of fibers are formed. The pith, which consists of thin-walled parenchyma at maturity, contains occasional nests of sclereids, although such nests are fewer than in the pith of Stenopadus or Stomatochaeta. Despite differences in size among species (e.g. large in C. mirabilis, very small in C. rupicola) no essential differences in anatomy could be detected. The differences among the three above genera seem related to habit features. The prominence of sclerenchyma in Stenopadus may be correlated with its arboreal habit. The presence of a sclerified epidermis on Stomatochaeta stems is a diagnostic feature, while the limited sclerenchyma in pith and cortex, to- gether with prominent cork formation, are characteristics which occur only in Chimantaea. Cortical sclereids have been reported for Stifftia (Vuillemin 1884), a genus which appears on other grounds to be closely related to Stenopadus. Elsewhere in Compositae, the only report of cortical sclereids appears to be that of Quentin (1911) in Baccharis. It is worthy of note that nests of sclereids in pith and cortex of stems and peduncles are very prominent in a Hawaiian mem- ber of Gochnatinae, Hesperomannia (Carlquist 1957e). The genus Quelchia (fig. 5), although a member of Gochnatinae on account of its actinomorphic corollas, is described here in connection with Gerberinae because of its stem structure. In Quelchia, as well as in Gongylolepis (fig. 4), diffuse sclereids are absent in cortex and pith. In both of these genera, how- ever, sclerification of parenchyma cells in areas between the conspicuous bundle eaps occurs. Mature pith of Gongylolepis consists of lignified parenchyma cells, although a few thick-walled sclereids were seen in pith of G. benthamiana. Pith of Quelchia conforms to the pattern seen in Gongylolepis. In all species of Gongylolepis examined, a bundle cap is present on cortical leaf traces. A few such fibers may also be present on the xylem pole of these traces. In Quelchia, however, fsbers occur only on the phloem pole of such bundles. Achnopogon and Dwidaea are like Quelchia in this respect ; they have relatively few interfascicular sclereids, however. In contrast, Neblinaea possesses the same prominent cylinder of sclerenchyma around the bundles as shown for Gongylolepis in figure 4. More- over, occasional diffuse sclereids are present in the cortex of Neblinaea. and cortical leaf traces are encased in a complete cylinder of fibers, in much the same manner as those of Glossarion (fie. 7). Pith in Achnopoaon, Duidaea, and Quel- chia is like that in Glossarion. Notable in the stem structure of Glossarion is the Fig. 4. Gongylolepis benthamiana, Maguire 33225; portion of transection of stem with secondary growth, cortex at left. Fic. 5. Quelchia cardonae, Wurdack 565; portion of cortex - transection, epidermis at left. Fig. 6. Gongylolepis huachamacari subsp. neblinensis, Maguire, Wurdack § Bunting 37353; portion of tangential section of cortex, showing laticiferous cells (shaded). Fic. 7. Glossarion rhodanthum, Maguire, Wurdack §& Bunting 37190; portion of transection of stem with little secondary growth; xylem and pith separated at right. Wend AEH My % Ah Me y ys) ein At 695 ARTY HR SATE 200, 2, DOs e eteas wessee ef 050 By SS ans esp One!!! . 9 oH? 0 GS fA wiles {0 Yopriee BSG (i qt DiPrc Veate sof pees o 8oss a2 WEG YAO: (oy eons oto Bia (oA Siii) 6 op O i As 6 ) y e a, wO0 G @) Oe OxXxe 0 0 A fe OLE} Oe, LY 162 MEMOIRS ‘OF THE NEW YORK BOTANICAL GARDEN [vou. 10 high degree of cortex sclerification, exceeding even that in Stenopadus. Because of this combination of features, it is difficult to ally Glossarion with either the gvongylolepoid or stenopadoid type. Differences among the various genera in respect to periderm origin and abundance are quite evident. Information from approximately comparable stems may be summarized as follows, periderm origin being noted where material permitted : Stenopadus: moderate, origin in third or fourth hypodermal layer Stomatochaeta: scanty, origin in third or fourth hypodermal layer Chimantaea: abundant, successive periderms in young stems Quelchia: moderate, origin in second or third hypodermal layer Gongylolepis: moderate, origin in second hypodermal layer Achnopogon: prominent, origin in first or second hypodermal layer Duidaea: prominent, origin in first or second hypodermal layer Neblinaea: seanty, origin in first hypodermal layer Glossarion: prominent, origin in first hypodermal layer Subepidermal origin of cork, such as occurs in these taxa, has been found in various Compositae by Douliot (1889), who did, however, record epidermal cork origin in the mutisioid Barnadesia. In none of the stems studied was there any indication of endodermis forma- tion, either in the form of Casparian strips, such as occur in Heliantheae (Carl- quist 1957d), or unevenly sclerified cells, such as occur in Barnadesia. Laticifers. The occurrence of laticifers in Compositae belonging to tribes other than Cichorieae was not generally realized until Col’s (1904) demonstra- tion of their occurrence in Cynareae, Vernonieae, and Arctotideae. It is of inter- est that two genera included by Hoffman (1890) in Mutisieae, Berardia and Warionia, were found to contain laticifers. Col concluded on the basis of lati- cifer occurrence and other characters that these two genera belonged in Cynar- eae, a treatment followed by subsequent.authors, such as Cronquist (1955). The discovery that stems of liquid-preserved material of Gongylolepis huachamacari subsp. neblinensis and Neblinaea promontorium contain laticiferous cells was therefore of considerable interest to the writer. The nature of the laticiferous structures in these two species enabled intrepretation of similar structures in other genera. Laticiferous cells in the cortex of Gongylolepis hwachamacari subsp. neblinensis (fig. 6), are often somewhat larger and more rounded than neighboring cells. While some laticiferous cells, by virtue of their extent, might suggest an articulat- ed series from which the intervening walls had been resorbed, no such walls were encountered, and the occurrence of groups of cells with intact walls suggests that at least predominantly we are concerned with individual cells, some of which enlarge and invade intercellular spaces. Other laticiferous cells, however, merely remain rounded. That the contents of these cells was indeed latex was confirmed by the milky appearance of cells en masse, as well as under higher magnification. Moreover, contents of these cells resisted safranin, a stain which is readily absorbed by resinous materials. That some laticiferous cells, particularly in outer portions of the cortex of G. huachamacari subsp. neblinensis, did contain resins was suggested by the presence of larger globules, which did stain deeply with safranin. Such cells proved to be present not only in cortex, but also in pith, leaves (fig. 21), and other parts of Gongylolepis and its allies, as well as Quelchia. Identification of laticiferous cells in herbarium material was based partly on their greater size, as compared to surrounding cells, and partly on 1958] ANATOMY OF GUAYANA MUTISIEAE—PART II 163 the fact that the clearing techniques employed did not completely remove cell conterits, which thus stained in whole mounts and sections of cleared parts. The taxa in which these were located with reasonable certainty are as follows: Gongylolepis huachamacari subsp. neblinensis: cortex, pith, leaf G. benthamiana: cortex, pith (fig. 4), involucral bracts G. fruticosa: cortex, leaf Duidaea pinifolia: cortex, leaf, achene Neblinaea promontorium: cortex, pith, peduncle Quelchia cardonae: cortex, leaf, achene Although laticiferous cells may occur more widely in the plant body than is indi- cated for the above taxa, their presence in the genera listed is of considerable interest. With the exception of Achnopogon, in which their presence could not be definitely established, laticiferous cells are present in all the Guayana genera of the Gerberinae. The presence, in addition, of laticiferous cells in Quelchia is noteworthy in view of its systematic position, and it may be that Quelchia is closer to Gerberinae than Stenopadus, Stomatochaeta, Chimantaea, and Glos- sarion, which apparently do not possess laticiferous cells. Beside the interest inherent in a new record of laticifers for the tribe Mutisi- eae, the fact that they are of the type found by Col in Arctotidae, Cynareae, and Vernonieae is of possible significance. Col, however, records anastamosing articulated laticifers in addition in Gazania of Arctotidae and in the subtribe Carlinae of Cynareae. The writer considers, however, that Col’s designation (p. 169) of individual laticiferous cells as ‘formes imparfaites .. . . témoinage d’une évolution extensive ou régressive’’ is open to doubt. It may be that in such individual laticiferous cells, singly or in groups, we are witnessing a primi- tive, rather than a degenerate form of the well-developed laticifers character- istic of Cichorieae. Secretory canals. The term ‘‘secretory canal’’ here refers to the structures sometimes designated as ‘‘oil canals’’ or ‘‘resin canals’? by various authors. Where present, they are always an intercellular formation, thus differing from laticifers, which are cellular in nature. Within the genera of this study no true secretory canals—in the sense of their discrete structure in Heliantheae—can be said to occur. In parenchyma or sclerenchyma of cortex and pith in Steno- padus and Stomatochaeta (figs. 1, 2), however, resins are abundantly secreted into intercellular spaces; these resins carbonize, giving a blackish color to the cortex and pith of these genera. Although such resin secretion is present even in leaves, involucral bracts, and achenes of these genera, in no case could a definite secretory canal be said to be present. It is of interest that secretion of resins into vessels and libriform fibers of the secondary xylem is also character- istic of these and other Gochnatinae (Carlquist 1957b). By contrast, none of the Gerberinae studied showed this type of resin secre- tion. It may be that secretion of resinous materials into the laticiferous cells replaces deposition in intercellular spaces in these genera. True secretory canals occur in some, although they are absent in many genera of Mutisieae. Van Tieghem (1872) has reported them in the root of Stifftia chrysantha, Gerbera schimperi, and others. He notes their presence in stem and leaf of Ainsliaea acerifolia, Hyalis spartiodes, Lycoseris mexicana, Mutisia retusa, Nassauvia digitata, and Polyachrus villosus. Recently, Cabrera (1951) has dem- onstrated their presence in Aphyllocladus denticulatus, A. ephedroides, and Plazia. It is of interest, therefore, that secretory canals occur in all subtribes 164 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN (vou. 10 of Mutisieae. Because of this fact and the infrequency of such canals in the tribe (see Solereder, 1908, for a list of Mutisieae in which they do not occur), the phylogenetic interpretation is not clear. The presence of unorganized inter- cellular deposition of resinous products may represent a stage before develop- ment of organized canals, or it may represent a substitution for such canals following their loss phylogenetically. Node and Petiole. The following numbers of leaf gaps were found for the genera of Guayana Mutisieae: Stenopadus: 5 Gongylolepis: approximately 7 Stomatochaeta: 3 Achnopogon: 3 Chimantaea: 5 Duidaea: 3 Quelchia: 3 Neblinaea: 5 Glossarion: 5 Considering that this group is fairly closely-knit, these divergences among genera within close alliances is noteworthy, and may probably best be explained on the basis of habit and leaf morphology in each genus. For example, the wide, sheathing bases of Gongylolepis leaves might be expected to be connected with a multilacunar condition, whereas the narrow or needle-like leaves of Duidaea seem correlated with a trilacunar situation. The phylogenetic significance of these differences, however. is difficult to interpret in the lack of adequate data for nodal conditions for the family. Both trilacunar and multilacunar nodes have been reported for the family (Sinnott 1914). It may be noteworthy in this connection that putatively primitive members of Heliantheae possess multi- lacunar nodes (Carlquist 1957d). In genera with sessile leaves (Stomatochaeta, Achnopogon, Duidaea) little branching of traces takes place in the cortex subtending the leaf, or in the leaf base. In the petiolate taxa, however, the traces branch into a simple are of numerous bundles in the petiole base. Lateral bundles of this are may be branched in their extent in the cortex, shortly above their departure from the vascular cylinder. The disposition of fibers around leaf traces in the cortex, mentioned earlier, continues in the petiole, and becomes altered into bundle sheaths, considered below. Bs Il. LEAF The Guayana Mutisieae are notable for excessive complication in leaf anatomy. The only other member of the tribe for which leaf anatomy has been described in detail, Hesperomannia (Carlquist 1957¢c) is much simpler by comparison. Hesperomannia has a mesophyll undifferentiated into palisade and spongy tissue, and is otherwise unspecialized except in its sunken biseriate hairs. | Leaves of the genus Stenopadus (figs. 8-11) show examples of complicated leaf structure, as well as demonstrating considerable specific differentiation in anatomical details. Leaves in this genus all have a thick cuticle on both surfaces and relatively thick leaves, but vary in: relative thickness of mesophyll; presence or absence of a hypodermis on adaxial or abaxial surface, and thickness of it; number of palisade and spongy layers, if differentiated. These features may be conveniently summarized in tabular form (figures for spongy and palisade are approximate). % 166 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [voL. 10 number of number of number of number of layers in layers in palisade spongy adaxial abaxial parenchyma parenchyma hypodermis hypodermis layers layers S.campestris (fig. 9) 2 1-2 3 7 S.connellii (fig. 11) 3 0 2 19 S. cucullatus (fig. 10) 1-2 0 2 iat | S. huachamacari 2 0 9 15 S. kunhardtii 2-3 1-2 3 0% | S. obconicus (fig. 8) 1-2 0 0 14 S. sericeus 3 0 2. 12 S. stipitatus 0 0 3 11 Variation among the species is found in other characters. Only one species, S. cucullatus, has bundle sheath extensions, these being present on a few larger veins. In S. obconicus and S. huachamacari, a few mesophyll cells are converted into thin-walled sclereids, mostly in layers near either epidermis. Bundle sheaths in some species (S. kunhardti, S. campestris, S. cucullatus, and 8S. stipitatus) have abundant fibers, particularly on larger veins. Where fibers are well devel- oped in the bundle sheath, they often intervene between phloem and xylem of the bundle itself. Some species are characterized by a sclerification of the lower epidermis (8. cucullatus, and, to a lesser extent, S. sericeus and S. connellir). Uniseriate hairs are retained on the lower surface of S. sericeus leaves, although the remaining species lose them early in ontogny, this variation being reminis- cent of the situation in Hesperomannia (Carlquist 1957¢). Each of the species of Stenopadus investigated, then, has a distinctive com- bination of characters, so that identification of a species by means of leaf anatomy would be quite feasible. The specializations in leaf anatomy have probably occurred quite independently of other changes in this genus, and while the different types of anatomy described are diagnostic, they do not fall readily into a phylogenetic series or into sections of the genus. However, the conversion of the lower epidermis into sclereids and the presence of diffuse sclereids in the mesophyll would seem to be advanced characters. Reference to leaves of other Mutisieae, subtribe Gochnatinae, aids both in showing that the patterns found in Stenopadus are relatively coherent, despite their diyersity, and in demonstrating that both less specialized and more ad- vanced conditions exist in other genera. In Stifftia chrysantha (fig. 29), which is considered related to Stenopadus, no true hypodermis is present, the meso- phyll is only about 9 cell layers in width, with poor differentiation into palisade and sponey tissue, and bundle sheaths are strongly fibrous. Bundle sheaths of larger veins are often in contact with the adaxial epidermis. In another genus of Gochnatinae, Anastraphia. (fig. 28), a more specialized. condition is seen. Although the leaf is similar in thickness to that of Stifftia, a prominant hypo- dermis is present. Moreover, the very conspicuous fibrous bundle sheaths form extensions reaching from hypodermis to lower epidermis in all but the smaller veins. Between veins, sunken pockets, covered with hairs and containing stomata projecting above the epidermis, are formed. Such stomata have been reportd by Solereder for the closely-related genus Gochnatia. A definite division into palisade and spongy tissue occurs. In Anastraphia, then, some of the features of Stenopadus and Stifftia leaves are present in an exaggerated form. 1958 | ANATOMY OF GUAYANA MUTISIEAE—PART II 167 Returning to Guayana Gochnatinae, the genera Stomatochaeta and Chiman- taea present striking comparisons with Stenopadus in leaf anatomy. Both of these genera, although they vary in leaf shape, have characteristic anatomical features which are correlated with the gross morphology and texture of the leaves. Some of the characteristics of Stomatochaeta leaves are shown diagrammati- cally in S. cymbifolia, figure 12. The epidermis of this species is unsclerified. A prominent hypodermis encircles the leaf, consisting of bands of thick-walled sclereids at the margins, and thin-walled sclereids in a single layer near both surfaces. Around the veins, thin-walled sclereids form a bundle sheath, and within this, on larger veins, a sheath of fibers is also present. No bundle sheath ioe oe Fies. 12-16. Sections of Stomatochaeta leaves. Fic. 12. S. cymbifolia, Steyermark & Wur- dack 508 ; diagram of transection; symbols explained in text. Fig. 13. S. condensata, Maguire 40603 ; portion of leaf transection, adaxial face above. Fic. 14. S. cylindrica, Maguire 32782; transection of leaf margin, adaxial face of leaf at right. Fic. 15. Same, face view of epi- dermis. - Fic. 16. Same, showing upper epidermis in transection; note abortive hair. ‘ 168 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vout. 10 extensions are formed, however . The mesophyll consists wholly of spongy paren- echyma cells, some of which become lheghtly sclerified at maturity. Although the leaves of S. condensata (fig. 13) are broader, the anatomy is much the same as that of S. cymbifolia. No hypodermis is present, however, and the subepidermal sclerenchyma in the leaf margins is not so extensive as that of S. eymbifolia. In Stomatochaeta cylindrica (fig. 14), a sclerified epidermis is present on the lower surface, but on the adaxial surface a sclerified hypodermis and non-sclerified epidermis are formed. In addition, a division into four layers of loosely arranged palisade and nine layers of spongy parenchyma occurs in the mesophyll of 8S. cylindrica leaves. The sclerenchyma of the margin (fig. 14, below) is approxi- mately as in S. condensata. Details of the epidemis of S. cylindrica are shown in figures 15 and 16. In figure 15, a surface view of the abaxial epidermis shows the thick-walled sclereids, often irregular in outline, which compose the entire epidermis with the exception of the guard cells and a single circle of cells surrounding each stoma. Guard cells are slightly external to adjacent epidermal cells, and the portions of guard cells which overlap epidermal cells are shown by dotted lines. Figure 16 illustrates the extremely thick cuticle which occurs on both surfaces of leaves of all species of Stomatochaeta studied. Also note- worthy is the occurrence of sunken hairs; these trichomes consist of a tier of rather short, wide cells, and do not project above the surface in mature leaves. Such abortive hairs, sunken into cuticle, are apparently characteristic of several taxa investigated, since they were observed also in Stenopadus and Quelchia. The genus Chimantaea (figs. 17-19) shows a highly individual series of fea- tures. Characters common to all species studied include an unsclerified adaxial epidermis with extremely thick cuticle, a prominent hypodermis at least two cells in thickness, subhypodermal sclereids with extremely thick walls scattered among the palisade cells, presence of spongy parenchyma, lower epidermis with relatively thin cuticle and exserted stomata, and an abundant development of uniseriate thick-walled trichomes on the lower surface. Chimantaea mirabilis (figs. 17, 18) has narrow leaves with inrolled margins. The epidermal cells are large, usually tapered toward the outside surface. The hypodermis, which is 2—4 cells in thickness, is composed of sclereids: About 3 layers of palisade and three of spongy parenchyma are present. Occasional in the palisade are enormous thick-walled sclereids, which span the distance between hypodermis and spongy parenchyma. The bundle sheath of veins consists of thin-walled sclereids, and although these cells are continuous with the hypo- dermis, which is composed of similar cells, such contact is not made between vein sheaths and the lower epidermis of larger veins. Consequently, sheath exten- sions cannot be said to be present. Although the leaves of Chimantaea mirabilis show the most extreme speciali- zation in their inrolled margins and exceptional developments of sclerenchyma, the other species also show a degree of expression of such features. Chimantaea- similis (fig 19), C. rupicola, and C. lanocaulis agree in having small epidermal cells, which are flat rather than tapered. The hypodermis in these species is formed of sclerenchyma much more thin-walled than in C. mirabilis, although it is wider, consisting of 4 or 5 cell-layers. Thick-walled mesophyll sclereids are abundant, usually occurring in groups. Such sclereids are not as long as the width of the palidsade, however. In all three species, approximately five layers of pali- sade and five of spongy parenchyma are present. The foliar anatomy of Chiman- 1958 | ANATOMY OF GUAYANA MUTISIEAE—PART II 169 BESAE 7 AN Ss, Says f par, S ve —— =n —, WT 17 8: F 17-20 yes, adaxial e ab Fig 17. Ch t bil Steyermark § Wurdack 821; diagram (symbols explained text). Fig. 18. Same, port f leaf transection. Fic. 19. C. similis, Steyermark 605; port fi) 0 € IG. : , , Quelchia cardonae, Steyermark § Wurdack 565; portiton of leaf transection. \ 170 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 taea is in certain respects like that figured by Marloth (The Flora of South Africa, Vol. 3, Pt. 2, 1932) for another genus of Gochnatinae, Oldenburgia. Chimantaea differs in its thick sclereid-bearing palisade; the palisade of Olden- burgia is narrow and composed of thin-walled cells. The similarities between the two genera probably represent parallel adjustments to xeric situations. The other genus of Guayana Gochnatinae, Quelchia (fig. 20) is distinguish- able in its leaf anatomy from the preceding by virtue of the fact that thin- walled sclereids form bundle sheath extensions on the larger veins. The two species studied, Q. bracteata and Q. cardonae, have extensions composed of cells identical in shape and wall thickness to those of the hypodermis. The adaxial hypodermis is approximately two cells in thickness, and a single layer of cells composes the abaxial hypodermis. Cells appearing to be laticiferous by virtue of contents and large size occur in the adaxial hypodermis. A division into three or four layers of palisade and about ten of spongy parenchyma occurs. A promi- nent cuticle is present on both upper and lower epidermis. Among Gerberinae, the genus Gongylolepis (figs. 21, 22) is least specialized in its leaf anatomy. From liquid-preserved material of G. hwachamacari subsp. neblinensis (fig. 21), the distribution of laticiferous cells could be determined. Cells containing latex occur in both upper and lower epidermis, in all layers of the adaxial hypodermis, and in spongy and palisade parenchyma. Gongylo- lepis huachamacari subsp. neblinensis exhibits an advance over other species of the genus in possessing a hypodermis which consists of a single layer of small thick-walled cells, together with two layers of parenchyma only slightly more thick-walled than palisade cells. A bundle sheath extension is present on most veins. Within the bundle sheath, an exceptional development of fibers is seen; these often extend to both epidermises, and intervene between the phloem and xylem of a vein. Approximately two layers of palisade and seven of spongy parenchyma are characteristic of G. huachamacari subsp. neblinensis. In .G. martiana (fig. 22), a condition more typical of the genus may be observed. One or two layers of hypodermis are present on the adaxial surface. The bundle sheath is highly fibrous, and extends to both surfaces in larger veins, forming extensions. Approximately three palisade and seven spongy layers are present. Among species of Gongylolepis investigated which are not illustrated, G. huachamacari and G. bracteata differ from G. martiana in having a thicker leaf, with 4 or 5 hypodermal, three palisade, and about 12 spongy layers. The bundle sheath and sheath extensions consist of cells like those of the hypodermis, rather than being composed of fibers. Fibers are, however, present immediately adja- cent to the bundles. The spongy cells immediately adjacent to the lower epi- dermis are often thickened, suggesting the development of a hypodermis on the lower surface. Gongylolepis fruticosa, a rather specialized species, shows notable differences in that the leaves are isolateral, with 3-5 layers of hypodermis on each surface, separated by 7 or 8 layers of spongy parenchyma. Bundle sheaths and sheath extensions are as in G. huachamacari. “ The genus Neblinaea, not illustrated, easily fits into the ranve of variation found in Gongylolepis. The description above of G. huachamacari could be applied to the only species, N. promontorium, except for the narrower hypo- demis (2 cell-layers), palisade (2-8 cells), and spongy tissue (about 5 cells). Achnopogon and Duidaea, by contrast, show remarkable specializations. The single species of Achnopogon (fig. 24) has a leaf anatomy basically similar to 1958 | ANATOMY OF GUAYANA MUTISIEAE—PART II 171 that of species of Gongylolepis. Bundle sheath extensions are more conspicuous, howeyer. Although the three layers of palisade are loosely arranged, they can be differentiated from approximately seven spongy layers. No hypodermis is present on the abaxial surface between the bundle sheath extensions. Similar features are seen in leaves of Duidaea (figs. 25-27) ; they are how- ceco? TA BS0RNQA0O99BOOSeSO oes er jal BOL OOO Spee 2 aa ra, e\/ | yA La a the Se Sao) Coe rent (PART. ole saa ot; Ds: LAD Vw vee es ney aa Nee LENS 1) es Sg oy) 72 OR Sy) @ SSO) be 2607 300 Ca ead Se cs >, TOS ere t\ ASG eS : Oe ROSY Figs. 21-24. Transections of leaves, adaxial face above. Fic. 21. Gongylolepis huachama- cari subsp. neblinensis, Maguire, Wurdack §& Bunting 37353; \aticiferous cells shaded. Fic. 22. G. martiana, Schultes 5742. Fic. 23. Glossarion rhodanthum, Maguire, Wurdack § Bunting 37190. Fic. 24. Achnopogon virgatus, Steyermark §& Wurdack 742. ‘, 172 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 ever, modified in accordance with the narrow, in some species needle-like, shape. In each of the three species studied, a prominent extension is present on the bundle sheath of the midvein, although lateral veins lack them. Little differen- tiation into palisade and spongy parenchyma can be discerned, except in D. rubiceps (fig. 27), where two layers of palisade and seven of spongy tissue may be recognized. Thick cuticle is present on both surfaces of the leaves of all spe- cies. Stomata in Duidaea leaves are confined to the lower surface on either side SES tO BN Sen ; K WHE ay Zw 9S ER ett; H) Of Figs. 25-29. Transections of leaves, adaxial face above. Fic. 25. Duidaea tatei, Steyer- ~ mark 58204; diagram (symbols explained in text). Fic. 26. D. pinifolia, Steyermark 58143; transection entire leaf. Fig. 27. D. rubriceps, Tate 1042; portion of transection including mid- vein. Fig. 28. Anastraphia cowellii, Howard 5098 (A); portion of transection. Fig. 29. Stiff- tia chrysantha, Trinidad Bot. Gard. 1330. (GH); portion of transection. 1958 | ANATOMY OF GUAYANA MUTISIEAE—PART II 173 of the midvein sheath extension (see figs. 25, 26). Fibers are abundantly present around the midvein in this genus, although lacking in the lateral veins; lateral veins are, however, jacketed by cells like those of the midvein sheath extension. The three species of Duwidaea may be very easily separated on the basis of hypo- dermal sclerenchyma. Duidaea tater (fig. 25) has a prominent adaxial hypo- dermis, composed of sclereids, which extends around the margins, where it is broader. Duidaea pinifolia (fig. 26), on the contrary, is entirely lacking in such sclereids. Duwidaea rubriceps (fig. 27) possesses a hypodermis a single layer thick (except at the margins, where it is two cells in thickness) which completely encircles the leaf. The leaf of Glossarion (fig. 23) combines features of many of the taxa de- scribed above, although it cannot be referred to any particular genus. A thick cuticle is present on both upper and lower epidermis. A hypodermis, consisting of a single layer of small isodiametric sclereids, occurs on the adaxial face. One feature not seen in any of the other genera is the elongate shape of the thin- walled sclereids composing the adaxial half of bundle sheath extensions. The lower half of the sheath extension consists of isodiametric thin-walled scleren- chmya, which also forms sheaths on smaller veins not possessing extensions. No hypodermis is present adjacent to the lower epidermis; numerous uniseriate hairs, not shown in the portion illustrated, characteristically occur on the lower epidermis. In summary, excellent generic distinctions are present among most of the eenera, such as Stomatochaeta, Chimantaeca, Duidaea, and Glossarion. The leaves of Stenopadus could be distinguished from those of Gongylolepis by means of the presence of laticiferous cells in the latter. The leaves of Stenopadus, in addition, tend to be much thicker than those of Gongylolepis. Quelchia leaves resemble those of both Stenopadus and Gongylolepis, although they are closer to the latter in possession of bundle sheath extensions (absent from Stenopadus except for S. cucullatus) and laticiferous cells. Leaves of Achnopogon and Neblinaea are within the range of variation found in Gongylolepis. The complex structure of leaves in Guayana Mutisieae, their abundant scler- enchyma or thick-walled parenchyma, massive cuticles, and possession of bundle sheath extensions are features usually associated with plants of habitats dry during at least part of a year. The leaf structure of such species as Chimantaea mirabilis is what one would expect to find in a rather xeric environment, in fact. One notable feature which is found in most of the genera is a development to some extent of a hypodermis, a structure which occurs (together with sheath extension) in such a distant mutisioid as Nowelia. The absence of hypodermis in such genera as Stifftia and Hesperomannia (Carlquist 1957¢) does indicate that although it is a widespread phenomenon in the tribe, it is not a universal one.’ Although investigations of leaf anatomy of other mutisioids are apparently lacking, comparisons can be made with other Compositae. Hypodermis has been reported in species of Celmisia (Astereae) and in Senecio munroi by Solereder (1908), and in Veronia luschnathiana by Grimm (1904). Bundle sheath exten- sions are apparently not infrequent in the family, as is indicated by reports of their occurrence in various species by Betts (1920), Shields (1951) and Carl- quist (1957d). In none of the genera studied by these authors, however, does ‘ 174 MEMOIRS. OF THE NEW YORK BOTANICAL GARDEN [vou. 10 there appear to be such a conspicuous representation of sclerenchyma; the excep- tional development of such cells in leaves, as well as in other parts, appears to be characteristic of Mutisieae. III. INVOLUCRE Structure of involucral bracts within a single species varies from outer to inner bracts. Likewise, the tip of a bract may differ in anatomy from the base. Fias. 30-34. Diagrams of transections of outer involucral bracts, adaxial face above (sym. bols explained in text). Fic. 30. Stenopadus connellii, Wurdack 34265. Fic. 31. Stenopadus cucullatus, Maguire 35120. Fic. 32. Stifftia chrysantha, Trin. Bot. Gard. 1330 (GH);. Fig. 33. Wunderlichia mirabilis, Glaziou 2168G (UC). Fie. 34. Chimantaea similis, Steyermark & Wurdack 605. 1958] ANATOMY OF GUAYANA MUTISIEAE—PART II 175 Differences between base and apex are usually in the nature of a greater promi- nence of parenchyma and more numerous bundles near the base. A section mid- way along the length of a bract gives a reasonable idea of the nature of its strue- ture, and so such sections were employed in this study, unless otherwise noted. The variation from outer to inner bracts, however, may take various forms and must be considered when comparisons among taxa are made. Receptacular bracts, if present, represent the most extreme expression of the series from outer to inner. In the genus Stenopadus, variations in involucral bract structure occur chiefly in the relative amount and distribution of sclerenchyma. Figures 30 and 31 show diagrammatic transections of outer involucral bracts of S. connellii and S. cucullatus respectively, and portions of inner bracts are shown in cellular detail in figure 35 for S. campestris and in figure 38 for S. cucullatus. Comparison of outer and inner bracts for a single species, S. cucullatus (figs. 31, 38), shows differences in thickness of sclerenchyma on each face of the bract and in number of bundles present. In both, however, the epidermis and several underlying layers on both sides of the bract consist of thick-walled sclereids. In the vascular bundles, fibers or isodiametric sclereids surround most of the bun- dle; they do not, however, separate xylem from phloem. There are few fiber groups in the mesophyll of bracts which are not directly related to bundles. In 8. connellu (fig. 30) the distribution of sclerenchyma is quite different. One or two layers on the inner face of the bract are unsclerified, although a thick band of sclereids is present below the surface on this face of the bract. The outer face of the bract is not sclerified, although individual sclereids or nests of sclereids occur throughout the mesophyll of the bract, unlike the condition in S. cucullatus. Sclerification is conspicuous in association with the bundles: the xylem pole of each strand is often completely surrounded, with the phloem lying on the abaxial face of this fibro-sclereid sheath. In addition, a fibrous bundle cap is formed adjacent to the phloem. Inner bracts of S. connellii, however, are hke those of S. cucullatus in that both surfaces of the bract are sclerenchyma- tous, and although fibers occur in association with the bundles, no sclerenchyma other than this is found in the mesophyll, which consists of thin-walled par- enchyma. Stenopadus campestris (fig. 35) does not agree with either of the two above species in that although the outer bracts are like those of S. connellii, the struc- ture of the inner bracts is different from that of S. connellii. Sclerenchyma, rather than being present on both faces, is more frequent toward the inner face, and occurs in the form of nests or individual sclereids. Sclereids are much less frequent toward the outer face of the bract. Among other species of Stenopadus investigated, S. kunhardtii and S. stipitatus agree with the types of bracts which occur in S. campestris; S. obconicus has inner bracts like those of S. cucullatus. A difference was noted above between the excessive fiber development in S. connellii and the fibrous sheaths of bundles in S. cucullatus outer bracts, in which an area of fibers (horseshoe-shaped in transection) often surrounds the bundle, but does not separate xylem from phloem. Among the other species of Stenopadus studied, S. obconicus resembles S. cucullatus in this respect, whereas S. campestris, S. kunhardtii, and S. stipitatus are like S. connellii in such char- acteristics of bundles in outer bracts. Bundles of inner bracts tend to be alike in all the species, with a more or less complete cylindrical sheath of fibers sur- .Y 176 MEMOIRS.OF THE NEW YORK BOTANICAL GARDEN [vou. 10 rounding vascular elements but not separating xylem from phloem. The pres- ence of carbonized resin deposits, among both parenchyma and sclerenchyma cells, was noted in bracts of several species of Stenopadus. Stomatochaeta (fig. 37) possesses bract structure similar to that of the inner bracts of Stenopadus cucullatus. As demonstrated by the bract of Stomatochaeta cylindrica shown in figure 37, however, a marked difference occurs between the the large-celled sclerenchyma on the outer face of the bract and the much smaller sclereids on the inner face. Inner bracts of Stomatochaeta are similar, except that sclereids on the outer face of the bract grade into the thin-walled Bats ‘s Ij 0 9 b BOLIC Eatoae es a 3 Se) O48: 6 &, eo GS x 6 4 A y 2 Sa ae rt) Se Deg, aS ISIN iS fepart LS velo et xj Sega ae (SS J NS one 5 U, Figs. 35-39. Portions of transections of involucral bracts, adaxial face above. Fic. 35. Stenopadus campestris, Maguire 35573; inner bract. Fic. 36. Chimantaea similis; Steyermark § Wurdack 605; inner bract. Fic, 37. Stomatochaeta cylindrica, Maguire 32782; outer bract. Fig. 38. Stenopadus -cucullatus, Maguire 35120; inner bract. Fic. 39. Quelchia cardonae, Steyermark & Wurdack 565; inner bract. 1958 | ANATOMY OF GUAYANA MUTISIEAE—PART II 177 parenchyma gradually, so that cells with walls of intermediate thickness may be found beneath the abaxial sclerenchyma. No nests of sclereids, however, are present in the mesophyll of bracts, although bundles are very prominently encased by fibers. In outer bracts, xylem and phloem of bundles are frequently separated by such fibers. The bracts of Chimantaea (figs. 34, 36) are distinctive in having more abun- dant sclerenchyma development than any of the preceding. This sclerenchyma, however, is not disposed as in Stenopadus or Stomatochaeta, since neither fibrous bundle sheaths nor nests of sclereids are present. Instead, the entire bract, both inner and outer, is converted into sclerenchyma except for a small band (in outer bracts, fig. 34) or occasional pockets (in inner bracts, fig. 36) of thin- walled parenchyma on which the vascular bundles are located. In their extremely woody structure, these bracts surpass those of other genera in this study. Involueral bracts of Quelchia (fig. 39), although they are thinner, have a combination of characters which does not match those of the above genera. The adaxial epidermis, and parts of one layer beneath it, are sclerified. as is the abaxial epidermis, but the remainder of the bract is thin-walled parenchyma. No fibrous bundle sheaths are present. The outer bracts have the structure shown for an inner bract of Q. cardonae in figure 39 except that the abaxial epidermis is unsclerified. Comparisons with other Gochnatinae seem pertinent here, both for demon- strating the relative similarity of bract structure, despite certain differences, in the four above genera, and for showing the diversity to be found in putatively closely-related members of this tribe. The only other mutisioid for which infor- mation on bract structure is available is Hesperomannia (Carlquist 1957c). so that two other genera, which are probably not distantly related to Guayana Gochnatinae, are described here. Stifftia (fig. 32) is notably different in that its bracts, although they have a texture like that of Chimantaea bracts, have an entirely different development of sclerenchyma. The adaxial epidermis may be sclerified. All other sclerenchyma in the bract, however, is in the form of dis- crete fiber bundles, and no isodiametric sclereids are present in the mesophyll. Some of these fiber bundles contain vascular components, although vascular tissue in bracts of Stifftia is at a minimum. Where xylem is present, it occurs in the center of one of these fiber strands, with the phloem at the outside of the strand separated from it by numerous fibers. The involucral bracts of Wunderlichia (fig. 33) are notable in that a promi- nent band of sclerenchyma covers the inner face with the exception of the epi- dermis. In this respect it resembles the bract of Stenovadus connellii shown in figure 30. In contrast to S. connellii, however. no sclerenchyma of any kind is associated with the vascular bundles. Grouns of parenchyma cells may develop into mesophyll sclereids which are much larger and more thin-walled than those - of the adaxial sclerenchyma. A quite different pattern is illustrated by the involucral bracts of Hespero- mannia (Carlquist 1957c). In this genus, no sclerenchymatous sheath is present on bundles. A thin band of sclerenchyma is present on the inner face of the bract including the epidermis, and there is a rather prominent development of fibers toward the outer face. The latter sclerenchyma may be outside of or intercalated between the bundles. A similar pattern oceurs in Anastraphia cowellii (Howard 5098, GH) with the exception that in addition to the isolated ‘ 178 MEMOIRS’ OF THE NEW YORK BOTANICAL GARDEN [vou. 10 band of fibers near the outside of the bract (interior to which are the bundles), the outer epidermis is sclerified. Two to three layers of sclerenchyma are present on the inner face of the bract; thus, three distinctive bands of sclerenchyma are present. Vascular bundles are mostly without associated fibers. In all char- acters except the abaxial epidermis, bract structure in Anastraphia cowelli is close to that of Hesperomanmna. SUITOIN ee) \S PEDO HOON OY ODGOKY Fias. 40-44. Portions of transections of involucral bracts, adaxial face above. Fia. 40. Achnopogon virgatus, Steyermark §& Wurdack 742. Fie. 41. Glossarion rhodanthum, Maguire, Wurdack & Bunting 37190; outer bract. Fic. 42. Duidaea rubriceps, Tate 1042; inner bract. Fig. 43. Gongylolepis huachamacari, Maguire et al. 30262; inner bract. Fic. 44. Gongylolepis martiana, Schultes 5742 (GH); outer bract. 1958] ANATOMY OF GUAYANA MUTISIEAE—PART II 179 In Guayana genera of the subtribe Gerberinae, a contrasting condition to all of the above can be seen in the genus Gongylolepis (figs. 43, 44). Sclerenchyma in Gongylolepis is notably absent. In the outermost bracts of G. martiana (fig. 44), the adaxial epidermis is sclerified and 2-4 underlying cell layers have mod- erately thick walls, but no other sclerification occurs in the ground tissue of the bract. Fibers are absent around vascular bundles. Cells in the abaxial half of the bract have contents which suggest the laticiferous-resinous content of cells elsewhere in the plant. In G. huachamacari (fig. 43), inner bracts show no sclerenchyma, and even outer bracts lack it altogether. The very fleshy texture of bracts in Gongylolepis, then, is in marked contrast with the bracts of genera described above. The other genera of Guayana Gerberinae show clear relationship to Gongy- lolepis in bract structure, although they are somewhat more specialized. The condition in Achnopogon (fig. 40) is virtually identical with that described for Gongylolepis martianus, although the bracts are thinner. Thick-walled paren- chyma cells occur beneath the sclerified inner epidermis. In inner bracts, no sclerenchyma is present except in the inner epidermis. No fibers or true scler- enchyma surround the vascular bundles. In Duidaea (fig. 42), precisely the same pattern is seen. As shown for D. rubriceps, however, no sclerenchyma is present, only parenchyma with moder- ately thick lignified walls, forming a single hypodermal layer. Such cells may also be present around the vascular bundles, particularly in the basal portions of the bracts. One or two layers of fibers were found to be present on the adaxial face of bracts of D. tatei. Bracts of Neblinaea are not unlike those of Duwidaea or Achnopogon except that they contain a single layer of fibers on both faces of the bract. In addition, fibers also encase the vascular bundles. Within the Guayana Gerberinae, then, some differentiation alone generic lines has taken place in respect to bract struc- ture, although the unity of the group is evident. This unity is evident if comparison is made with another genus placed in Gerberinae (Mutisinae of Hoffman, 1880), Nowelia. In outer bracts of NV. insignis (Rock 11714, A) a thick band of sclerenchyma. including the epidermis. forms the inner face. The abaxial epidermis likewise is sclerified. Bundles are sheathed with fibers, although additional strands consisting wholly of fibers are present. Inner bracts differ only in that the adaxial epidermis alone is sclerified, the fibrous bundle sheaths being as in the outer bracts. The genus Glossarion is anomalous in respect to bract structure, just as it is in other features. The bract shown in figure 41 shows an unsclerified outer epi- dermis with thick cuticle overlying 2-3 ecell-layers of thin-walled sclerenchyma, which contrasts with the markedly sclerified cells on the inner face of the bract. Fibers encase the vascular elements. Within the thin-walled parenchyma cells - of the mesophyll, there is wide variation in cell size. In lower portions of bracts of Glossarion, the parenchyma is more uniform, and all cells of the bract except the vascular bundles and the sclerenchyma on the inner face could be classified as thick-walled parenchyma. While the bract structure of Glossarion is not un- like that of several genera considered here. particularly Stenopadus, it cannot be clearly referred to any of them. Receptacular Bracts. Only a small minority of Mutisieae possess recentacular bracts. Because two genera included in this study (Stenopadus and Stomato- 180 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN ) Lh ‘oa SES < AR QQ ANY S WIRQAAY AS WS SSN Ss (WJ \ 49 v (oy Te iy SOY SS SN \ \ le ‘ | r fi —S—— 1960] BOTANY OF THE GUAYANA HIGHLAND—PART IV 19 This species differs from the closely related P. wurdackii in its completely lepidote leaf-blades and subsessile flowers. Brocchinia delicatula L. B. Smith, sp. nov. Fig. 6. Florifera 20-35 em alta; caule erecto, 3 em vel altiore, 8 mm diametro, basi- bus vetustis atris foliorum dense vestito; foliis apice caulis rosulatis, ad 16 cm longis, dissite adpresseque lepidotis, vaginis latissime ovatis, 1 em longis, atro- castaneis, sublucidis, laminis linearibus, acuminatis, 4 mm latis, planis, dense nervatis, pallide viridibus, integris; scapo erecto, gracillimo, glabro; scapi bracteis erectis, remotis, parvis, anguste triangularibus, dissite lepidotis; inflor- escentia laxe bipinnatim paniculata, 5-13 em longa, glabra; bracteis primaris eis Scapi similibus, quam basibus sterilibus nudis ramorum brevioribus; ramis divergentibus, 3-5 em longis, gracillimis, laxe paucifloris; bracteis florigeris ovatis, acutis, 2.5 mm longis, membranaceis; floribus divergentibus, breviter pedicellatis; sepalis oblongis, obtusis, 2.5 mm longis, membranaceis, straminels ; petalis oblongis, obtusis, 3.5 mm longis, haud unguiculatis, albis; staminibus inclusis, filamentis cum sepalis petalisque brevissime coalitis; ovario clavato, 5 mm longo, minime supero, stylis distinctis, gracilibus. VENEZUELA: Amazonas: Cerro de la Neblina, Rio Yatua, alt. 1100 m, locally abundant in rock crevices along Canon Grande east of Cumbre Camp, 24 Nov 1957, B. Maguire, J. J. Wurdack & C. K. Maguire 42207 (type). This species is by far the smallest and most delicate in Brocchinia and is easily distinguished by its habit alone. It belongs with the species with promi- nently nerved leaf-blades and glabrous inflorescences but does not appear closely related to any of them. Navia diffusa I. B. Smith, sp. nov. Fig. 7. Florifera ad 3 m alta; foliis interioribus solum cognitis, ad 4 dm longis, sub- dense serrulatis, supra glabris, vaginis late ovatis, ca. 4 cm longis, brunneis, subtus lepidibus albidis orbicularibus adpressis subdense vestitis, laminis anguste triangularibus, nullo modo contractis, 27 mm latis, subtus obscure dissiteque lepidotis ; seapo 2 em diametro; scapi bracteis erectis, foliaceis, dense imbricatis ; inflorescentia laxissime 3-4-pinnatim paniculata, glabra; bracteis primarus late ovatis, acuminatis, quam basibus sterilibus ramorum brevioribus, infimis serru- latis; ramis ad 28 em longis; bracteis florigeris latissime ovatis, quam sepalis subduplo brevioribus; floribus patentibus, sessilibus; sepalis lberis, cochleari- imbricatis, late ovatis, obtusis, 4 mm longis; petalis Juvenilibus solum cognitis, inclusis; ovario supero. VENEZUELA: Amazonas: Cerro de la Neblina, Rio Yatua, alt. 1900 m, frequent on edge of north escarpment, 23 Dee 1957, B. Maguire, J. J. Wurdack §& C. K. Maguire 42537-E (type). Figs. 2, 3. Cottendorfia. Fig. 2. C. nubigena (Maguire, Wurdack §& Maguire 42151). a, base of leaf, X 0.5; b, branch of inflorescence, * 0.5; ¢, flower, XK 1; d, sepal, K 2. Fic. 3. C. savannensis (Maguire, Wurdack §& Maguire 42167). a, base of leaf, X 0.5; b, braneh, < 0.5; ce, flower, X 1; d, sepal, X 2. Fies. 4, 5. Pitcairnia. Fic. 4. P. wurdackii (Maguire, Wurdack & Maguire 41584). a, base of leaf, & 1; b, flower, & 1; ¢, sepal expanded, * 1; d, ovary, X 1; e, seed, X 5. Fig. 5. P. maguirei (Maguire, Wurdack § Keith 41556). a, base of leaf, & 1; b, flower, X 1; ¢, sepal expanded, X 1; d, ovary, X 1; e, seed, X 5. [von. 10 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN 1960} BOTANY OF THE GUAYANA HIGHLAND—PART IV 21 In- my key in the previous report (Mem. N. Y. Bot. Gard. 9: 295), this species runs down to the vicinity of N. brocchinioides but is quickly distin- guished by its short pinnately arranged spikes. Navia abysmophila L. B. Smith, sp. nov. Fig. 8. Acaulis vel breviter caulescens; caule ad 10 cm longo, pauciramoso, 5 mm diametro, mox nudo; foliis multis, apice caulis densissime aggregatis, ad 6 em longis, interioribus aream coloratam circa inflorescentiam haud formantibus, vaginis integris, infimis subtriangularibus, parvis, supremis suboblongis, ca. 1 em longis, laminis linearibus, acuminatis, 3.5 mm latis, basi haud contractis, subtus valde nervatis cretaceisque, mox utrinque glabris, spinis minutis subdense serrulatis ; inflorescentia subsessile, simplici, pauciflora, 5 mm diametro; bracteis exterioribus subfoliaceis sed valde reductis; bracteis florigeris late ellipticis, acutis mucronulatisque, 8 mm longis, omnino membranaceis, pallide brunneis, lepidibus minutis linearibus pallidis sparsissime vestitis ; sepalis oblongis, obtusis cucullatisque, 7 mm longis, posterioribus anguste alato-carinatis; petals albis; staminibus inclusis; ovario supero. VENEZUELA: Amazonas: Cerro de la Neblina, Rio Yatua, alt. 1100 m, locally frequent on rocks in bottom of Canon Grande south-southwest of Cumbre Camp, 24 Dee 1957, B. Maguire, J. J. Wurdack & C. K. Maguire 42471 (type). The species is also represented by number 42202 from east of Cumbre Camp. This novelty closely resembles Navia cucullata but differs in having the young leaves cretaceous beneath and the floral bracts acute and uniformly membranaceous. Navia affinis L. B. Smith, sp. nov. Fig. 9. - Caulescens; caule erecto, simplici, ad 12 em alto, 7 mm diametro, foliorum vaginis vetustis praeditis; foliis Jjuvenilibus terminalibus, subpetiolatis, ad 3 dm longis, apice obscure serrulato excepto integris, vaginis triangularibus, 34+ em longis, ex sicco rubentibus, margine villoso-lepidotis, laminis lineari-lanceolatis, acuminatis, 10 mm latis, marginibus paulo incrassatis; inflorescentia in centro foliorum sessili, ellipsoidea, 20 mm longa, 6 mm diametro, verisimiliter simplici; bracteis ellipticis, ca. 15 mm longis, sepala superantibus, subtenuibus, apice paulo incrassatis rubentibusque, dissite lepidotis, integri, laevibus; sepalis ob- longis, obtusis, 12 mm longis, tenuibus, 14 connatis; petalis albis, reflexis et stamina nudantibus; ovario supero. VENEZUELA: Amazonas: Cerro de la Neblina, Rio Yatua, alt. 1100 m, locally frequent on rocks in bottom of Cafion Grande south-southwest of Cumbre Camp, 24 Dee 1957, B. Ma- guire, J. J. Wurdack & C. K. Maguire 42470 (type). Fig. 6. Brocchinia delicatula (Maguire, Wurdack §& Maguire 42207). a, habit, X 0.5; b, branch of inflorescence, X 1; ¢, sepal and stamen, X 5; d, petal and stamen, X 5; e, ovary, X 5. Fies. 7-10. Navia. Fic. 7. N. diffusa (Maguire, Wurdack & Maguire 42537—-E). a, base of leaf, X 0.5: h. brareh of inflorescerce, K 0.5; ¢, flower, X 1: sepal expanded, X 5. Fig. 8. N. abysmophila (Maguire, Wurdack & Maguire 42471). a, habit, * 0.5; b, inflorescence, « 1; e, sepal, X 1. Fia. 9. N. affinis (Maguire, Wurdack §& Maguire 42470). a, leaf, & 0.5; b, flower, X 1. Fic. 10. N. parvula var. expansa (Maguire, Wurdack. § Maguire 42246). a, inflorescence, * 0.5; b, flower, * 1. 22 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Only further collections will prove whether the above is a distinct species or only a variety of N. subpetiolata. At present it can be said to differ by its nar- rower leaf-blades and smaller, more highly connate sepals. Navia parvula var. expansa IL. B. Smith, var. nov. Fig. 10. Caule elongato, pauciramoso; foliis ad 15 cm longis; floribus inflorescentiae pluribus; sepalo anteriori verisimiliter omnino abortivo; petalis erectis, anguste lanceolatis, 15 mm loneis, aureis; staminibus longe exsertis. VENEZUELA: Amazonas: Cerro de la Neblina, Rio Yatua, alt. 1200-1400 m, abundant in colonies on dry rock ledges in Canon Grande east of Cumbre Camp, 26 Nov 1957, B. Ma- guire, J. J. Wurdack g C. K. Maguire 42246 (type). Number 42502 from dry eliff faces in forest south-southwest of Cumbre Camp at alt. 1050-1100 m is the same variety, and 42536 from the same eliffs but higher and in a sheltered area is intermediate between the two varieties. The type of the species is its smallest form and occurs at the highest altitude, which indicates a correlation between shelter and size- of plant. The material from this latest expedition provides good flowers which were lacking in the type, and thereby discloses several unusual characters. The petals are narrow and erect and the stamens long-exserted, but most remarkable is the absence of any trace of an anterior sepal. Such a situation can be considered the final phase in a reduction evolution which began by its being covered by the other sepals. ANNONACEAE’ Guatteria longedecurrens R. E. Fr., sp. nov. Arbor ca. 18 m alta; ramuli novelli nigrescentes, pilis sparsis mox deciduis instructi; internodia 1-2 em longa. Foliorum petiolus ca. 5 mm longus, supra paulo canaliculatus; lamina 10-15 em longa, 3—5 em lata, ovato-lanceolata, basin versus longe acutata et in petiolum decurrens, rigida, apicem versus in cuspidem 1-1.5 cm longam obtusam sat sensim contracta, supra (exsiccata) sordide viridis, subtus olivacea; costa supra vix depressa, subtus valde prominens teres et rubescens; nervi laterales I tenues; reticulum venularum densissimum, utroque latere elevatum. Flos (unicus obvius) 2 em longe pedunculatus; pedicelli rigidi, olabri. Petala oblonga, apice rotundata, exteriora ca. 14 mm longa, 6 mm lata, interioragca. 15 mm longa, 7 mm lata. Stamina 1.5-2 mm longa, disco plano elabro. Monocarpia numerosa, immatura nigra, obovoidea, obtusa, 8-10 mm longa, 4-5 mm crassa, in stipitem 6-7 mm longum contracta. VENEZUELA: Merida, Pueblos del Sur, alt. 1600-2200 m, Bernardi 2255 (type NY, cotype S). This species agrees in most respects with G. glaberrima R. E. Fr. (known from Ecuador), which, however, even in early stages of development has quite elabrous branchlets and leaves. In G. longedecurrens the network of veins is very elevated and conspicuous, especially on the under side. It differs from G. glaberrima also in its broader, ovate-lanceolate leaves. Cxandra venezuelana R. E. Fr., sp. nov. Arbor alta; ramuli glabri; internodia 1-1.5 em longa. Foliorum lamina rigida, glaberrima, flavovirens, ovato-oblanceolata, basi rotundata vel brevissime TBy Robert E. Fries. 1960 | BOTANY OF THE GUAYANA HIGHLAND—PART IV 23 acutata, apice in cuspidem brevem acutam obtusiusculam sensim contracta, 8-13 em longa, 3-4 em lata; costa supra conspicue impressa, rubescens; nervi laterales et venularum reticulum densissinum utrinque conspicue prominentes. Flores in axillis foliorum delapsorum glomerati, fere sessiles; pedunculi brevissimi, post florationem ad 2 mm excrescentes, bracteolis minutis biseriatis retundatis aureo-sericels dense instructi. Sepala rotundato-ovata, obtusiuscula, extus dense breviterque sericea, ca. 2.5 mm longa et lata. Petala oblanceolata, 12-14 mm longa, 5-7 mm lata, ad marginem brevissime ciliata. Stamina numerosa, flavida, apice recurvata, 3-4 mm longa. Pistilla numerosa, clavato-cylindrica, dense tomentella, ovulo basali unico erecto instructa. (Fructus ignotus. ) VENEZUELA: Merida: municipio El Vigia, selva pluvial submontana, alt. 150-200 m, 5 Feb 1955, Bernardi 1923 [‘‘Exemplares medidos de mas de 50 Ms. con 7/3 del tronco libre de ramas’’] (type NY). This species is particularly characterized by the rigid leaves rounded at the base. In this respect it most closely approaches O. lewcodermis (Spruce) Warm., which also belongs to the Venezuelan flora. The new species proposed here is distinguished from the latter by a different nervation. The midrib is depressed on the upper side and the vein network is extremely close and elevated on both sides of the leaf. | Guatteria bernardi R. E. Fr., sp. nov. Arbor; ramuli novelli cortice nigro obtecti et pilis divaricatis sat densis vestiti; internodia ca. 1 em longa. Folorum petiolus 2—3 mm longus, hirsutus ; lamina 5-10 em longa, 1.5-2.5 em lata, membranaceo-rigida, exsiccata obscura, supra glabra, subtus pilis brevibus adpressis + dense vestita, anguste lanceolata, basin versus angustata et apice in cuspidem 1—1.5 ecm longum acutum econtracta. Flores in axillis fohorum solitarii vel raro bini; pedicelli rigidi, usque ad 1.5 cm longi, vulgo recurvati, pilis patentibus rufis vestiti et primo bractea foliacea ad 1 em longa mox caduea instructi. Sepala rotundata, acuta, mox valde recurvata, 4-5 mm longa. Petala erecta, tomentella, oblanceolata, demum 15—20 mm longa, marginibus valde recurvatis. Stamina 1 mm longa. Monocarpia numerosa, ovoidea, glabra, castanea, acuta, 9-10 mm longa, stipitibus rigidis, 4-7 mm longis sustenta. Semen ovoideum, glaberrimum, ca. 8 mm longum. VENEZUELA: Bolivar: Calzeta de la Botella, 14 Apr 1957, Bernardi 6534 (type NY, cotype S). This species agrees with some forms of G. schomburgkiana, but differs from that species especially in its stipitate monocarps; the flowers have longer pedicels and the petals also are notably longer. Furthermore, the new species here de- scribed resembles G. flavovirens R. BE. Fr. (also known from Venezuela) which, however, differs in its conspicuously prolonged leaf apices, shorter peduncles, and shorter petals. | Annona gigantophylla (R. E. Fr.) R. E. Fr., comb. nov. Xylopia gigantophylla R. E. Fr. Ark. Bot. II. 3: 605. 1956. A recent fruiting collection of this species (Maguire, Wurdack, & Magwire 42060-A, from the slope forest of Cerro de la Neblina) has revealed the typical fleshy fruit of the genus Annona. 24 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [von. 10 RUTACEAE® In the following account several taxa are described as new, but I have also included descriptions of older taxa which were not fully characterized originally or which are much better known now as a result of numerous recent collections, In citing the place of deposit of these collections, the reference to the New York Botanical Garden sheet of the Maguire et al. collections has been omitted, since it is certain that the first set of these will be found in the New York herbarium. Raveniopsis Gleason, Brittonia 3:166. Oct 1939. Shrubs or small trees, mostly with simple leaves. Inflorescence a simple raceme, or dischasial with two cincinnoid racemes and a terminal flower at the apex of the peduncle, or rarely the flower solitary; flowers pentamerous, the ealyx, corolla, and androecium variously zygomorphic. Sepals free or very shortly united basally, usually obviously foliaceous, generally considerably longer than wide, imbricate. Corolla tube cylindric, arcuate, flaring toward the apex in a spreading limb, the petals unequal. Fertile stamens two, the ligulate filaments more or less joined with those of the three or more staminodia in a filament tube, adherent to the corolla tube at the base or much higher, the anthers with saccate to flabellate appendages at their base. Disk cupular, usually completely enclosing the ovary, truncate to somewhat lobate. Stigma obliquely d-lobed, the style slender, the ovary 5-carpellate, the carpels free except in the styles; ovules 2 in each carpel. Fruit of 1-5 cocci, each one-seeded, dehiscing longitudinally, the exocarp separating from the whitish endocarp. Type species: Raveniopsis tomentosa Gleason, op. cit. 167. As Gleason indicated in his original publication of this genus, several differ- ences separate it from the allied genus, Ravenia. Before the present publication, Ravenna has been considered to occur both in the lowlands and on the summits of the table-mountains of Guayana, but the species of the latter region are herein transferred to Raventopsis. The principal, technical characteristic that separates the two genera, as interpreted here, is the presence of basal appendages on the two fertile anthers of Ravenmopsis; but there are at least two other tendencies: (1) in Raven the flowers are solitary or borne in few-flowered, subumbellate clusters or racemes, but in the alhed genus the flowers are usually rather closely arranged in simple or dichotomous racemes or spikes; (2) in Ravenia the sepals are about as long as wide and not obviously foliose, in Raveniopsis the sepals are usually longer than wide and obviously foliose. 7 The species of Raveniopsis are found thus far exclusively on the summits of the tepuis in the Guayana Highland between 1000 and 2500 meters elevation ; the greater number of species occurs at about the 2000-meter level. Thus an altitudinal disjunction is correlated with the morphological distinctions and sup- ports the realignment of species presented here. . Key to the Species of Raveniopsis Leaves trifoliolate, chartaceous to thinly coriaceous, never rigid. ae _ — . . Leaves unifoliolate or simple, rigid. bo ° Inflorescence a simple raceme; larger sepals narrowly lanceolate, about 1.5 mm wide. Pubescence simple. 1. R. trifoliolata. 8By Richard S. Cowan. 1960] BOTANY OF THE GUAYANA HIGHLAND—PART IV 29 2. Inflorescence dichasial, consisting of two cincinnoid racemes and a terminal flower at the apex of the peduncle (cf. Fig. 10d); larger sepals broadly ovate or ovate- oblong, 1.5-5 mm wide. Hairs compound, dendroid with the branches closely aggre- gated on the basal part of the main axis, the latter long-attenuate in a more or less rigid point. 2. R. stelligera. 3. Hairs simple 4, 3. Hairs compound, stellate or appearing so. 10. 4. Leaves linear, strongly revolute. Flowers about 6 mm long, the corolla tube nearly straight. 8. R. linearis. 4. Leaves elliptic, oval, or ovate. Flowers longer than 6 mm, the corolla usually ob- viously arcuate. 5. 5. Inflorescence spicate, the calyx thick, densely pilose-velutinous. Leaves and stems densely velutinous. 11. R. nubicola. 5. Inflorescence dichasial (cf. Fig. 10d), consisting of two cincinnoid racemes and a terminal flower at the apex of the peduncle, or the inflorescence reduced to a solitary flower. 6. 6. Flowers solitary. Vegetative parts hispid, the leaves ovate to broadly so. 7. R. paruana. 6. Flowers in dichasia (ef. Fig. 10d). Vegetative parts not hispid, the leaves elliptic, oval, or ovate. iF 7. Leaves pilose on the undersurfaces. Corolla white; sepals elliptic-oblong. Fertile anthers lanceolate, mucronate by a prolongation of the connective about 1 mm long. 6. R. abyssicola. 7. Leaves strigulose, sericeous, or tomentose on undersurfaces. Corolla pale lilac, pink, or red; sepals ovate, lanceolate, or oblanceolate to elliptic. Fertile anthers oblong, not mucronate. 8. 8. Leaves strigulose only on primary veins beneath. Style pilosulose; corolla red. 3. R. ruellioides. 8. Leaves sericeous or tomentose on undersurfaces, the latter obscured by the pub- escence. Style glabrous; corolla pale lilae or pink. 9. 9. Larger sepals broadly elliptic, 3-3.5 mm long; corolla tube 5.5 mm long. Leaves sericeous beneath. 4. R. sericea. 9. Larger sepals oblanceolate to narrowly elliptic, 5.5 mm long; corolla tube 9 mm long. Leaves usually tomentose beneath. 5. R. fraterna. 10. Pubescence buff-canescent, densely aggregated on most parts, the lower leaflet surfaces completely obscured, the hairs dendroid. Leaves strongly convex-revolute, not mucronate. Inflorescence spicate. 9. R. tomentosa, 10. Pubescence red-brown, not obscuring completely the lower leaf surfaces, the hairs stellate, stipitate. Leaves about plane, the midrib often extended beyond the blade in a short, rigid, blunt mucro. Inflorescence dichasial, consisting of two ecincinnoid spikes and a terminal flower at apex of the pedunele (ef. Fig. 10d). 10. R. peduncularis. 1, Raveniopsis trifoliolata Cowan, sp. nov. Fig. 19. Arbusculae 1-2 m altae, ramulis rubidis et minute strigulosis. Folia digitato- trifoliolata (interdum unifoliolata) ; petioli 27-40 mm longi, graciles, minute strigulosi, foliolorum lateralium petiolulis 1.5-3 mm longis, minute strigulosis, folioli terminalis petiolulis 5-10 mm longis; foliolorum laminae chartaceae, ellipticae, 5-9 em longae, 3-4 em latae, acutae, foliolorum lateralium base in- aequilaterali, latere inferiore obtuso sed superiore attenuato-acuto, folioli terminalis base attenuato-cuneato, aequilaterali, supra glabrae, infra costa strigulosae sparse, costa salienti, venis primariis ca. 10-jugatis. Inflorescentiae axillares, solitariae vel binatae, unilaterale racemosae, 10-20 cm longae, ped- unculo 4-6.5 em longo, sparse puberulo, gracili, axe striguloso et puberulo, pedicellis 1-1.5 mm longis, minuto-strigulosis sparse. Sepala plus minusve chartacea, acuta, minute strigulosa sparse, duobus lanceolatis, 5.5—-7 mm longis, 1.5 mm latis, sepalis ceteris ca. triangularibus, 1-1.5 mm longis, 0.5-1 mm latis. Corollae tubus curvatus, glaber, 10.5-12 mm longus, infundibuliformis, lobo 10 mm longo, 3.5 mm lato, lobis ceteris ca. 5-8 mm longis, 1.5-3 mm latis. Stamina 5, duobus fertilibus, filamentis ligulatis, 15 mm longis, antheris 2.5 mm longis, 26 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN. [vou. 10 Fics. 11-17. Raveniopsis. Fie. 11. R. abyssicola (Maguire, Wurdack §& Maguire 42502). a, portion of branchlet, * 0.5; b, flower with one sepal removed, * 1.5; ¢, fertile anther, X< 5. bo 1960] BOTANY OF THE GUAYANA HIGHLAND—PART IV ad basem appendicibus flabellatis 1.3 mm longis ornatis ; staminodia usque 20 mm longa, ciliolata. Discus 0.9 mm profundus, usque 1.5 mm diametro, margine crenu- lata; stylus 10-15.5 mm iongus, glaber, ovarium sparse strigulosum. Fructorum cocci immaturum venosum, sparse strigulosum, mucronulatum. VENEZUELA: Amazonas: Cerro de la Neblina, Rio Yatua; locally abundant along Cafion Grande east of Cumbre Camp, alt. 1100 m; shrub 1-2 m; fis red; 24 Nov 1957, Bassett Maguire, John J. Wurdack & Celia K. Maguire 42217 (holotype, US no. 2267440). As the specific epithet emphasizes, this species is easily separable from most of the other species of the genus by the digitately trifoliolate leaves. The calyx is also very distinctive in shape and the glabrous corolla contrasts with the pukescent corolla of most of the other species in the genus. The simple hairs on many of the parts of R. trifoliolata separates it easily from its nearest relative, R. stelligera, which also has trifoliolate leaves but differs in having stellate pubescence, a cincinnoid-racemose inflorescence, and a different calyx. In the type collection are included a number of detached inflorescences which appear to be fasciated; instead of a simple raceme, which is characteristic of the species, the peduncle bears at its apex up to six racemose branches. 2. Raveniopsis steligera (Cowan) Cowan, comb. nov. Fig. 20. Galipea stelligera Cowan, Mem. N. Y. Bot. Gard. 8:121. Apr 1953. Shrub or small tree 2—5 m tall, the branchlets densely pubescent with cin- ~namon-colored, dendroid-stellate hairs. Leaves digitately trifoliolate, dendroid- stellate (except the intervein areas on the upper surfaces), the petioles 1.5—2 em. long, the petiolules of the lateral leaflets 1-2.5 mm long, that of the terminal one o—9 mm long;-leaflet blades chartaceous, elliptic to somewhat obovate or ob- lanceolate, the lateral leaflets smaller with the base inequilateral, the upper side attenuate-acute, the lower side rounded, 2—7 em long, 1—2.5 em wide, the apex acute or rounded and apiculate, the terminal leaflet larger, its base attenuate- acute, the apex acute, 4-7 cm long, 1—2.5 em wide, the lower surfaces of the leaflets densely dendroid-stellate-pubescent, much paler than the dark green upper surface, the latter sparingly to sparsely stellate-pubescent on the larger veins, the costa and primary veins impressed above, salient beneath. Inflorescence dendroid-stellate-pubescent, axillary or psuedoterminal, dichasial, the 2—-5-cm- long peduncle bearing at its apex one flower and two racemose branches, the bractlets about 1-3 mm long, subulate, caducous, the pedicels 0.5-1.5 mm long. Flowers pubescent with stellate and dendroid-stellate hairs, coral red to brick red, the sepals more or less unequal, oblong-oval to broadly ovate, obtuse or acute; corolla with a narrowly funnelform tube and erect limb, the tube slightly curved, 18—20.5 mm long, 6-7 mm in diameter, the adaxial corolla lobe triangu- lar-lanceolate to broadly ovate, the other lobes triangular to ovate-triangular, acute. Fertile anthers 2.5-3 mm long, oblong, apiculate, the appendages 1.5 mm long, ca. 1 mm wide, flabellate, the staminodia 18.5-20 mm long, ca. 1 mm wide, Fig. 12. R. sericea (Wurdack 34297). a, habit, * 0.5; b, flower, X 1.5; ¢, fertile anther, * 10. Fig. 13. R. fraterna (Steyermark § Wurdack 811). a, flower, X 1.5. Fig. 14. R. linearis (Tate 531). a, habit, X 0.5; b, transverse section of leaf, < 5; ¢, flower, X 3. Fic. 15. R. paruana (Cowan § Wurdack 31225). a, habit, X 0.5. Fic. 16. R. peduncularis (Maguire 32754). a, habit, X 0.5; b, flower, X 2. Fic. 17. R. nubicola (Maguire, Wurdack §& Magwire 42297). a, habit, X 0.5; b, calyx spread apart, «2. 28 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Fics. 18-21. Raveniopsis. Fic. 18. R. tomentosa (Cardona 2663). a, tip of fertile branch- let, X 0.5; b, dissected calyx, & 3; ¢, one hair, X 20; d, corolla, x 2. Fig. 19. R. trifoliolata 1960 | BOTANY OF THE GUAYANA HIGHLAND—PART IV 29 ligulate. Disk undulate marginally, ca. 0.8 mm deep; style glabrous, ca. 20 mm long, ovary densely hispidulous. Key to the Varieties of Raveniopsis stelligera 1. Mature calyx 3 mm long or shorter, the sepals plane, the larger ones not more than 2 mm wide, the smaller about 1.5 mm wide. Lateral leaflets acute. 2a. var. stelligera. 1. Mature calyx 4-5 mm long, some or all of the sepals reflexed at about the costa, the larger sepals broadly ovate, 4 mm wide, the smaller ones 2—2.5 mm wide. Lateral leaflets mostly rounded-obtuse, sometimes apiculate. 2b. var. plicata. 2a. Raveniopsis stelligera var. stelligera. Galipea stelligera Cowan, Mem. N. Y. Bot. Gard. 8:121 Apr 1953. Endemic to the summit of Cerro Duida at about 1650 m elevation, in scrub forest on the banks of streams (type collection, Maguire, Cowan & Wurdack 29653-A). 2b. Raveniopsis stelligera var. plicata Cowan, var. nov. A var. stelligera foliolis lateralibus rotundatis vel obtusis et apiculatis, 3 em longis vel brevioribus, calyce 5 mm longo, lobis reflexis in costa, lato-ovatis vel plus minusve oblongis, acutis, lobis 2 maioribus 4 mm latis, late ovatis, 3 min- oribus 2-2.5 mm latis, oblongis usque oblongo-lanceolatis, petalo adaxiali late ovato, ca. 3.5 mm longo et lato differt. VENEZUELA: Amazonas: Cerro Coro-Coro, Serrania Yutaje, Rio Manapiare, occasional in savannas and among rocks; alt. 1500 m; shrub 2—5 m high (sun form); fis coral red; 2 Mar 1953, Bassett Maguire § Celia K. Maguire 35440 (Holotype, US No. 2253691). Same data and collectors 35262, 35277, 35302 (paratypes). Endemic to the summit of Serrania Yutaje between 1300 and 2000 m elevation in woodland thickets and in open rocky savannas. As is indicated in the varietal key, these varieties are separated principally by the differences in the proportions of the calyx in addition to a geographic disjunction ; the plicate condition of the sepals is a striking difference separating var. plicata but it is not completely constant and hardly specific in importance. The difference in the shape of the adaxial petal in the two varieties is also worth noting, but it is mostly a matter of width. When I originally described this species I included it in the genus Galipea, but it cannot belong there, as I presently interpret that genus, because of the form of the corolla and calyx. The sepals in individual flowers of material of the typical variety are not so dissimilar as they are in var. plicata and, for that matter, in most of the species of Raveniopsis and at the time of the original description of this species I ignored the small differences. This species is one of three with the stellate type of pubescence; in this one and in R. tomentosa the hairs are dendroid-stellate. The branches of the hairs are more or less equally distributed along a main axis in that species; in R. stelligera the branches are rather closely aggregated on the lower part of the main axis and the greater part of the axis is naked of branches and drawn out (Maguire, Wurdack § Maguire 42217.) a, habit, X 0.5; b, calyx and gynoecium, X 2; ¢, corolla and androecium, < 2. Fie. 20. R. stelligera. a—e, var. stelligera; a, calyx, X 3; b, corolla, < 1; -c, one hair, X 20; d—g, var. plicata; d, habit, < 0.5; e, mature calyx, 3; -f, gynoecium, X< 3; g, corolla, X 1. Fig. 21. R. ruellioides. a—b, var. ruellioides (Maguire et al. 42447); a, habit, X 0.5; b, calyx, X 2; e-d, var. angusta (Maguire et al. 42189); ¢, habit, pee oe. dd, rcalyx, «K 2: 30 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN ~ [vou. 10 into a sharp, rather rigid point. This similarity in pubescence types and the form of the corolla limb suggests relationship with &. tomentosa, but in general habit R. stelligera is nearer R. trifoliolata, a species which is very different in the in- florescence, calyx, and type of pubescence. 3. Raveniopsis ruellioides (Oliver) Cowan, comb. noy. Ravenia ruellioides Oliver, Timehri 5:191. 1886. Key to Varieties of Raveniopsis ruellioides 1. Leaves oval to elliptic, about 2-3 times as long as wide, the base subobtuse to acute, not attenuate-cuneate. Sepals ovate to broadly ovate. da. var. ruellioides. 1. Leaves narrowly elliptic, about 3.5-4 times as long as wide, the base long- attenuate-cuneate. Sepals narrowly lanceolate. db. var. angusta. 3a. Raveniopsis ruellioides var. ruellioides. Fig. 21. Ravenia ruellioides Oliver, Timehri 5:191. 1886. Ravenia tatei Gleason, Bull. Torrey Club 58:376. Jun 1931. Ravenia ruellioides var. ptariana Steyermark, Fieldiana Bot. 28:273. Dee 1952. Widely distributed on the summits of sandstone tepuis from southernmost Venezuela to Mt. Roraima at 1500-2500 m elevation, in thickets and low scrub forest, especially alone streams. VENEZUELA: Amazonas: Cerro Sipapo, Maguire g& Politi 28095, 28211, 28271, 28459, 28667; Cerro Duida, alt. 1825-2000 m, Steyermark 58114 (NY), Tate 436 (NY); Cerro de la Neblina, alt. 1900-2100 m, Maguire et al. 42281 (US), 42447 (US). Bolivar: Mt. Roraima, alt. 2000-2500 m, Im Thurn 211 (isotype US), Irwin 428 (US), Pinkus 141 (G, NY, US), Steyermark 58715 (G, NY); Ilw-téput; Maguire’ 33459 (US); Cerro Apacara; Cardona 1544 (US); Auyan-tepui, Tate 1184 (NY, US); Ptari-tepui, Maguire § Wurdack 383891 (US), Steyermark 59511 (NY); Cerro Guaiquinima, Maguire 32941 (US), 32964 (US), 33088 (US); Chimantéa Massif, Steyermark 74893 (NY), Steyermark § Wurdack 393 (NY, US), Wurdack 32066, 34166 (NY, US). 3b. Raveniopsis ruellioides var. angusta Cowan, var. nov. Fig. 21. A var. ruellioide foliis anguste ellipticis, ca. 3.5-4-plo longioribus quam latis, ad basim longe attenuato-cuneatis; sepalis anguste lanceolatis differt. VENEZUELA: Amazonas: Cerro de la Neblina, rocky beaches along banks of Canon Grande east of Cumbre Camp, alt. 1100 m; locally frequent shrub 0.3-1.5 m tall; fis red; 24 Nov. 1957, Bassett Maguire, John J. Wurdack §& Celia K. Maguire 42189 (holotype, US No. BTN) Same data and collectors 42495 (paratype, US). The varietal key presents the characteristics that distinguish the two varieties. Among the materials of this highly variable species there are several forms, some of which have received names, but most of such variants are too inconstant to justify their recognition nomenclaturally. In the face of such a situation it may seem inadvisable to describe a new variety but it is sufficient- ly well separated morphologically and ecologically from the typical popula- tion that such a course cannot be satisfactorily avoided. f. ruellioides is perhaps nearest R. sericea and R. fraterna but it differs from these, as well as from other species of the genus, by its pilosulose style, by the distribution of the pubescence on the leaves, and by the color of the corolla. 4. Raveniopsis sericea Cowan, sp. nov. Fig. 12. Arbusculae 0.5-2 m altae, ramulis dense flavo-sericeis. Folia simplicia, rigido-coriacea, petiolis 3-6 mm longis, flavo-sericeis; laminae 28-40 mm longae, 17-30 mm latae; late ellipticae, elliptico-ovatae vel ovatae, ad apicem ob- tusae, ad basem rotundo-obtusae, pilosae sed @labrescentes supra (pilis suberectis ~ 1960] BOTANY OF THE GUAYANA HIGHLAND—PART IV 31 usque ad subappressos), infra dense flavo-sericeae, ad marginem valde revolutae, costa venis primariisque conspicue impressis supra, infra valde salientibus. In- florescentiae cymiformes, terminales, flavo-sericeae, pedunculo 20-35 mm longo, flavo-sericeo, pedunculi apice unum florem terminalem et duos ramos racemi- formes ca. 10-20 mm longos ferenti. Calyx dense sericeus, sepalis 2, late ellipticis, acutis, 3-3.5 mm longis, 2-3 mm latis, sepalis ceteris linearibus, 2.5-3 mm longis, 0.5—0.7 mm latis. Corolla sericea extus, intus in fauce minute lanulosa, tubulari- infundibuliformis, tubo 5.5 mm longo, lobis inaequalibus, ca. 3 mm longis, 1-2 mm latis, anguste oblongis et anguste ovafis. Stamina fertilia 2, filamentis ligulatis, 0.5 mm. longis, glabris, antheris oblongis, 1.2 mm longis, flabellato- appendiculatis, staminodiis 4, linearibus, pilosis. Stylus 2.6 mm longus, glaber; ovarium quinquelobatum, disco cupulari immerso. Fructorum cocci, sericel, oblique ovati, ca. 4 mm longi, 3 mm lati; semen papillatum, nigrum, ca. reni- forme, 2 mm longum. | VENEZUELA: Bolivar: northwest cumbres, Churi-tepui (Muru-tepui), frequent in lower part of lower cumbre, alt. 2100-2200 m; shrub 0.5-2 m; corolla very light pink, with a pink stripe centrally outside on each lobe; 24 Jan 1953, John J. Wurdack 34180 (holotype, NY); same locality and collector, alt. 2300-2500 m, 34297 (paratype, NY, US). The nearest relative of this species is R. fraterna, which differs in having a longer corolla tube and sepals of a different shape, and the pubescence on the under surfaces of the leaves is different. R. paruana is another, more distant, relative; it differs in having bright orange-red flowers which are borne singly at the tips of the sprawling branches; the type and distribution of the pubescence are also distinctive. 5. Raveniopsis fraterna Cowan, sp. nov. Fig. 13. Arbusculae 1-1.5 m altae, ramulis dense tomentosis. Folia simplicia, rigida, petiolis 2-3 mm longis, tomentosis; laminae 20-28 mm longae, 10-17 mm latae, ellipticae usque ad ovales, obtusae ad apicem basemque, longo-pilosae sed mox slabrescentes supra, infra dense tomentosae, ad marginem valde revolutae. costa venis primariisque valdissime impressis supra, infra valde salientibus. Inflor- escentiae terminales, racemosae, pedunculo 6-20 mm longo, tomentoso. Calyx tomentosus usque ad sericeo-tomentosum, 2 sepalis oblanceolatis usque ellipticis, acutis, 5.5 mm longis, 3 mm latis, sepalis ceteris lanceolato-linearibus, 5 mm longis, 0.8 mm latis. Corolla tomentella extus, intus in fauce puberula, tubulari- infundibuliformis, tubo 9 mm longo, lobis inaequalibus, 3-4.5 mm longis, 24 mm latis, oblongis vel suborbicularibus. Stamina fertilia 2, antheris oblongis, 1.5 mm longis, appendicibus flabellatis, 0.5 mm longis, staminodiis 5, basilariter pilosulis, ca. 2 mm longis, linearibus. Stylus 8 mm longus, glaber; ovarium quinauelo- batum, in disco cupulari inclusum. Fructorum cocci pilosa ad apicem, 4.5 mm longi, ca. oblongi; semen echinato-papillatum, fuscum, oblongum 3 mm longum. VENEZUELA: Bolivar: Torono-tepui, Chimanté Massif, summit at edge of escarpment in and among zanjones, alt. 2165-2180 m; shrub 1-1.2 m, locally frequent; lvs spreading, sub- coriaceous, revolute, dark green above with suleate veins, grayish-buff below with elevated veins; calyx dull green; corolla tube whitish with dull lilac, the lobes white inside and flushed pale lilac outside; 9 Feb 1955, J. A. Steyermark §& J. J. Wurdack 646 (holotype, NY); Cen- tral Section, Chimanta Massif, open forest on laterite deposit above swamp following east branch of headwaters of Rio Tirica, alt. 2150-2200 m, 12 Feb 1955, Steyermark & Wurdack 811 (paratype, NY, US); Chimanté Massif, along edge of ridge at summit of east-central portion of summit of Apacara-tepui, alt. 2450-2500 m, 21-22 Jun 1953, Steyermark 75905 (paratype, NY). - 32 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [von. 10 R. fraterna is wost nearly related to R. sericea; the differently shaped sepals, the longer corolla tube, and the pubescence on the under surfaces of the leaves of Rk. fraterna are distinguishing characteristics. In this latter characteristic there is not complete uniformity; the type collection has the leaves distinctly and densely tomentose, the hairs rather flexuous, but the paratype collections show this feature less strikingly and in some examples the hairs are even more or less appressed. The flower parts are, however, quite consistently unlike in the two species. 6. Raveniopsis abyssicola Cowan, sp. nov. Fig. 11. 3 Arbuscula 2-3 m alta, ramulis pilosis, rubicundis. Petioli graciles, subap- presso-strigosi, 5-14 mm longi, folliis simplicibus; foliorum laminae coriaceae, discoloratae, infra pallidiores, ellipticae, ad basem attenuato-cuneatae, apicaliter acutae, ad marginem angusto-revolutae et ciliatae, 6.5-9 em longae, 2—3.5 cm latae, supra glabrae (pilosa costa excepta), infra pilosae, costa et venulis planis supra, infra salientibus, venulis ca. 20-jJugatis, in vena marginali conjugentibus. Inflorescentiae axillares, dichasiales, 6.5-7 em longae, axé piloso, pedunculo 4 em longo, bracteis caducis, 1.5—-2.5 mm longis, anguste ellipticis; pedicelli 1.5-2.5 mm longi, strigosi. Sepala glabra intus, subappresso-strigosa extus, acuta, duo ex- teriora 6-8 mm longa, 2—2.5 mm lata, anguste elliptico-oblonga, duo interiora 4 mm longa, 1.25 mm lata, sepalo quinto 4 mm longo, 0.5 mm lato, lineari. Corolla alba, sericea valde et subabrupte curvata, tubo 11-12 mm longo, 1.5 mm diametro ad basem, apicaliter ca. 4 mm diametro, lobis 5-7 mm longis, 1.25—-2.5 mm latis, obtuso-acutis. Stamina fertilia 2 adaxialia, filamentis 7.5 mm longis, ligulatis, sericeis ad apicem, tubo cum staminodiis conjunctis, antheris anguste oblongis, sericeis, ca. 1.5 mm longis et 0.7 mm latis, connectivo in subulatum mucronem ca. 1 mm longo producto, antherarum basi biappendiculata, appendicibus parvis, ca. 0.8—-0.6 mm longis, staminodiis ligulatis, ca. 13 mm longis, plus minusve villoso sericeis. Discus cupularis, haud erassus, 0.7 mm altus; ovarium ca. 0.9 mm altum, sparse pilosulum. Fructorum cocci cornuti, 6-7 mm longi, 3 mm lati, ovati, mucronati, sparse pilosuli; semina subreniformia, 3-3.5 mm longa, 1.5-2 mm lata, fulva, verruculosa ad minute muricata. VENEZUELA: Amazonas: Cerro de la Neblina, occasional in bottom of Cafion Grande ssw of Cumbre Camp, alt. 1050-1100 m; shrub 2-3 m; fis white; fruit green; 25 Dec 1957, Bassett Maguire, John J. Wurdack § Celia K. Maguire 42504 (holotype, US No. 2267451). The Specific epithet for this remarkable species is chosen to emphasize its habitat preference—the bottom of the nearly mile-deep cafion on Cerro de la Neblina. Several striking distinctions combine to separate R. abyssicola from all the other species of the genus: (1) the corolla is decidedly more zygomorphic than that of any of the others; (2) the fertile anthers have very small, hardly notice- able appendages at the base and unlike those of any of the other species are tipped by the elongate connective. There are also abundant vegetative character- istics to distinguish the species. 7. Raveniopsis paruana (Cowan) Cowan, comb. nov. Fig. 15. Ravenia paruana Cowan, Mem. N. Y. Bot. Gard. 8:122. Apr 1953. VENEZUELA: Amazonas: summit of Cerro Part, Serrania Parti, Feb 1951, Cowan ¢ Wurdack 31285 (type collection). Apparently endemic to Cerro Part, growing in thickets in rocky scrub- savanna. 1960] BOTANY OF THE GUAYANA HIGHLAND—PART IV 33 The solitary flowers of this species separate it from all others in the genus; the strongly hispid character of the vegetative parts is also distinctive. 8. Raveniopsis linearis (Gleason) Cowan, comb. noy. Fig. 14. Ravenia linearis Gleason, Bull. Torrey Club 58:375. Jun 1931. VENEZUELA: Summit of Mount Duida, alt. 1050-1657 m, Dee 1929, G. H. H. Tate 531 (type collection); same locality, Steyermark 58133 (NY), 58212 (NY). Endemic to Cerro Duida, at 1050-1675 m elevation, along the banks of streams. This is by far the smallest-flowered species, and the size and shape of the leaves furnish further distinguishing characteristics. 9. Raveniopsis tomentosa Gleason, Brittonia 3:167. Oct 1939. Fig. 18. VENEZUELA: Bolivar: Auydén-tepui, alt. 2200 m, G. H. H. Tate 1362 (type collection, NY); same locality, Cardona 2663 (NY, US). Apparently endemic to Auyan-tepui, Venezuela. The pubescence of this species is dendroid-stellate with the branches more or less evenly distributed along the main axis of the hair. In respect to pub- escence at least, R. peduncularis is the closest relative; in that species the central axis of the hairs is not developed and a stalked, many-armed, star-shaped form results. The form of the inflorescence and flower parts, as well as vegetative differences, separates the two species. 10. Raveniopsis peduncularis Pittier & Lasser, Bol. Soc. Venez. Ci. Nat. 9:121. 1944. Fig. 16. Shrub 1-3 m tall, the branchlets densely and darkly rusty-brown-stellate- pubescent. Leaves simple, rigid, strongly discolorous, the petioles stout, 4-10 mm long, densely brown-stellate-pubescent, the blades (3.5—)5.5-8.5 em long, (1.5—) 24.5 em wide, elliptic to ovate to narrowly ovate, obtuse basally, the apex acute, often with a mucro to 2 mm long, the upper surface at first densely grayish- stellate-pubescent but soon glabrescent and then nitid and distinctly green when fresh, the lower surface densely rusty-stellate-pubescent, the costa im- pressed above but strongly salient beneath, the venules plane and subobscure above, prominulous beneath. Inflorescences spicate, dichasial, densely rusty- stellate-pubescent, the peduncle arising in the upper axils, 2.5-4 em long, the two unilateral branches to 5 em long. Flowers orange-red, sessile, rusty-stellate- pubescent externally, the calyx densely so; sepals strongly unequal, obtuse, more or less joined very shortly at the base, the two largest 4 mm long, 1.5 mm wide, oblong-lanceolate, the remaining three sepals 2-3 mm long, ca. 1 mm wide, glabrous within except at the apices; corolla tube slightly arcuate, 8-11 mm long, villose within, the limb strongly unequal, the upper lip entire, 7 mm long, 2 mm wide, lanceolate, the lower lip 4-lobed, the lobes lanceolate-oblong, ca. 3.5 mm long, 1 mm wide. Filaments of the two fertile stamens 12 mm long, villose, the fertile anthers oblong, ca. 2 mm long, glabrous, the staminodia 10-13 mm long, villose. Disk shallowly cupulate, 0.6 mm deep, 1.5 mm in diameter, completely enclosing the glabrous ovary, the style glabrous, 7 mm long, tapering apically to the digitate stigma. Cocci of the fruit rusty-red-stellate-pubescent, venose, apiculate, ca. 3 mm long, 2 mm wide, the seed minutely verruculose, reniform, ¢a. 2 mm long, 1 mm wide, dark brown. 34 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 VENEZUELA: Bolivar: Cerro Guaiquinima, upper Rio Paragua, Oct 1945, Cardona 948 (type collection, NY, US); same locality, alt. 1740-1760 m, Cardona 1089, Maguwire 32754 (US), 32815, 32854, 32944, 32985, 33069. Endemic to Cerro Guaiquinima; frequent at 1600-1800 m elevation on or just below the summit in scrub thickets or rocky, exposed areas and in scrub- savannas. In addition to the difference between the two types of stellate pubescence, this species differs from its nearest relative, R. tomentosa, in the type of in- florescence, as well as in many details of the flower and leaves. R. peduncularis is characterized by a dichasial inflorescence, the branches of which are unilater- ally spicate; the inflorescence of R. tomentosa is a simple, unilateral spike, obviously a reduction of the dichasial arrangement of its relative. 11. Raveniopsis nubicola Cowan, sp. nov. Fig. 17. Arbusculae 0.7-1.5 m altae, ramulis velutinis, foliis simplicibus. Petioli erassi, 7-12 mm longi, velutini; foliolorum laminae rigidae, valde discoloratae, ovales, obovatae, ellipticae vel late ellipticae, 37-83 mm longae, 20-38 mm latae, ad apicem obtusae et apiculatae, ad basim acutae, subobtusae vel obtusae, marginaliter valde revolutae, supra nitidae, glabrae (costa pilosa excepta), infra densissme luteo- vel cinnamomeo-tomentosae, costa Impressa supra, infra valde salienti, venulis subobscure impresso-reticulatis supra, infra salientibus, 12~15- jugatis. Inflorescentiae dense spicatae, interdum binatae, usque ad 6 em longas, densissime piloso-velutinae, pedunculo 10-17 mm longo, bracteis ca. 5 mm loneis, anguste lanceolatis, acutis, persistentibus ; flores sessiles, aurantiaco-rubri. Sepala crassa, densissime piloso-velutina, duobus maioribus 10 mm longis, 3 mm latis, obtusis, oblanceolatis, sepalis ceteris 6-7 mm longis, 1-1.5 mm latis, lanceolatis, acutis. Corolla curvata, extus pilosa in lineis, intus limbo glabro sed tubo tomen- tuloso, tubo ca. 10 mm longo, lobis 5.5-7.5 mm longis, plus minusve ellipticis, obtusis. Antherae 2.5 mm longae, anguste ellipticae, glabrae, dorsualiter sericeae, appendicibus 0.5 mm longis, filamentis 9 mm longis, anguste oblanceolatis, vil- losis. Discus 1 mm profundus, glaber, leviter crenulatus. Stylus 4 mm longus, elaber. Fructorum cocci usque ad 5, piloso-velutina, 4 mm longa, 3.5 mm lata, seminibus 3 mm longis, 2 mm latis, reniformibus, papillato-muricatis, nigris. VENEZUELA: Amazonas: Summit of Cerro de la Neblina, Rio Yatua, locally frequent in scrub forest 15-18 km sw of Cumbre Camp, alt. 2000 m; shrub 0.7-1.5 m; fils orange-red; 2 Dee 1957, Bassett Maguire, John J. Wurdack §& Celia K. Maguire 42297 (holotype US No. 2267443). This high-altitude representative of the genus is named for its seeming preference for the cloud-shrouded summit of Cerro de la Neblina. It is very surely most nearly related to R. tomentosa and R. peduncularis, more closely to the former than to the latter. R. nubicola differs in a number of characteristics : (1) its pubescence is simple, in contrast to the dendroid-stellate pubescence of its near relatives; (2) it has substantially longer petioles; (3) its sepais are differently shaped and also differ somewhat in size; (4) the corolla is red in the new species but white in its nearest relative; (5) the corolla lobes are longer and differently shaped in the new species. Myllanthus Cowan, gen. nov. Fig. 22. Arbor; folia opposita, trifoliolata. Inflorescentiae unilateraliter cincinnoideo- racemosae, cauliflorae vel ramuliflorae; flores pedicellati, hermaphroditi, sepalis 1960 | BOTANY OF THE GUAYANA HIGHLAND—PART IV 39 liberis, petalis valde imbricatis, tubo inaequali conjunctis; stamina fertilia 2, antheris basim appendiculatis, dorsualiter in connectivo valde papillato-glandu- latis, filamentis in tubo corolla cohaerentibus. Discus cupularis, quinquelobatus ; stigmata quinquelobata; ovarium quinquepartitum, ecarpellis solum stylo con- junctis. Fructorum cocci monosperma, capsularia. Type Species: Myllanthus maroana Cowan. Myllanthus is properly associated phylogenetically with two genera, Lubaria and Raputia. All three genera are characterized by zygomorphic corollas; in most of the species the corolla is more or less arcuate, and this characteristic is the basis of the generic name of the new genus (pvAAds, myllos—bent; av6os, anthos—flower). Lubaria is easily distinguished from the others by its free adaxial petal with the two fertile stamens on the adaxial side of the flower; the other two genera (Myllanthus and those species of Raputia with appendaged anthers and strongly arcuate corolla) have the abaxial petal more nearly free than the other petals (but distinctly joined with the others at least basally) and the two fertile stamens on the abaxial side of the flower. However, the basal appendages of the anthers are very different in the Raputia species, consisting of elongate, ligulate structures which are obviously the unmodified, sterile bases of the anther thecae. At least two species of Raputia have the arcuate corolla and the orientation of the androecium of Myllanthus, but in addition to the difference in the appendages, the calyx of the Raputia species is cupular with small teeth on the margin of the cup. Key to the Species of Myllanthus 1. Leaflets oblong, three times as long as wide. Inflorescence 3-4 em long, not obviously coiled. , 1. M. maroana. 1. Leaflets elliptic, less than three times as long as wide. Inflorescence 10-18 em long, obviously coiled. 2. 2. Leaflets with petiolules 1-2 em long; petioles about 8 em long. Flowers with pedicels 3-4 mm long. 2. M. neblinae. 2. Leaflets sessile; petioles 4 em long or shorter. Flowers sessile or subsessile. 3: M..uter. _ . Myllanthus maroana Cowan, sp. nov. Fig. 22, a—d. Arbor 3-6 m alta, ramulis minute malpighiaceo-strigulosis sed mox gla- brescentibus, foliis digitato-trifoliolatis. Petioli 10-12 em longi, glabri, canalicu- lati, laminae sessiles, oblanceolatae usque ad oblanceolato-oblongas, coriaceae, discoloratae, infra pallidiores, glabrae et nigro-punctatae, 25-45 em longae, 8-14 em latae, acutae apicaliter, ad basim attenuato-cuneatae, costa salienti, venulis planis supra, infra salientibus. Inflorescentiae cauliflorae, 3-4 cm longae, uni- lateraliter cincinnoideo-racemosae, axe et floribus minute malpighiaceo-strigulosis, pedicellis 4-6 mm longis. Sepala 3-3.5 mm longa et lata, orbicularia vel suborbi- eularia, ciliolata; petala villosa intus, inaequaliter connata, petalo abaxiali in tubo ca. 4 mm connato, ejus lobo 15 mm longo, obtuso, petalis ceteris in labia quadri- lobata, curvata ca. 11 mm longa connata, eorum lobis oblongis, leviter cucullatis, ad apicem rotundatis. Stamina 4 sed solum 2 fertilia, filamentis elongato- oblanceolatis, apicaliter subulatis, villosis; antherae fertiles oblongae, connectivo papillato-punctato, 5.5 mm longae, 1.5 mm latae, appendicibus late flabellatis, antherae apicem versus curvatis et papillato-glandularibus, ca. 2 mm longis et 4 mm latis. Discus carnosus, ca. 1 mm profundus; stylus 9 mm longus, glaber; ovarium glabrum, papillatum. Fructorum cocci oblongo-obovata, 17 mm longa, 10 mm. lata, ad apicem dehiscentia; semina ca. 12 mm longa, 8 mm lata, ovali- oblonga. 36 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN 1960] BOTANY OF THE GUAYANA HIGHLAND—PART IV 37 ° VENEZUELA: Amazonas: occasional in high forest along trail to Guzman Blanco 1-5 km se of Maroa, Rio Guainia, alt. 120-140 m; tree 3-8 m; eauliflorous; fis cream; 9 Oct 1957, B. Maguire, John J. Wurdack & W. M. Keith 41780 (holotype, US No. 2267414); Selva de Maroa, Rio Guainja, alt. 127 m, 17 Feb 1942, L. Williams 14392 (paratype). This species differs from M. neblinensis, its nearer relative, in size and shape of the leaflets, and in the length of inflorescence and the degree to which it coils; there are doubtless many good floral characteristics that separate the two species but the lack of mature flowers in M. neblinensis makes critical comparison im- possible. 2. Myllanthus neblinensis Cowan, sp. nov. Fig. 22, e-f. Arbor 5 m alta, ramulis petiolis petiolulisque minutissime malpighiaceo- strigulosis. Petioli 6.5—-7.5 em longi, petiolulis 1-2 em longis, haud alatis; laminae chartaceo-coriaceae, plus minusve concoloratae, glabrae, 16-21 em longae, 6-8 em latae, ellipticae, ad apicem acutae, ad basim attenuato-cuneatae, costa salienti, venulis planis et inconspicuis supra, infra salientibus. Inflorescentiae scor- pioideae, ca. 10 cm longae, axe et floribus minutissime malpighiaceo-strigulosis, pedunculo 20-27 mm longo, pedicellis 3-4 mm longis. Calyx ad basim truncatus, sepalis ciliolatis, rotundatis, 2.56 mm diametro, corolla androecium et gynoecium valde immaturis. Fructorum cocci oblonga, 10 mm longa, 7 mm lata, valde trans- verse costata, dense strigulosa. VENEZUELA: Amazonas: Cerro de la Neblina, Rio Yatua, in talus forest between Camps 3 & 4, alt: 900-1000 m; tree 5 m; buds greenish; 8 Nov 1957, Bassett Maguire, John J. Wurdack & Celia K. Maguire 42001 (holotype, US No. 2267436). Ordinarily this material would await description for further collections to be made but, since the slopes of Cerro de la Neblina are not likely to be visited again for a very long time, I have given this a name. It differs from its nearer relative, M. maroana, in leaf size and shape, and in characteristics of the in- florescence. The flowers are very immature but sufficiently developed to determine its affinity with this genus. 3. Myllanthus ulei (Krause) Cowan, comb. nov. Cusparia ulei Krause, Repert. Sp. Nov. 2:26. Jan 1906. BRAZIL: Amazonas: near Manaos, Ule 5382a (type). Lowlands near rivers in northwestern Brazil. Near Santa Izabel, Rio Negro, Ducke 339 (US); Manaos circa Cachoeira do Mindi, Ducke 20478 (US). There are perhaps other species of Cusparia which should be removed to other genera, but this one is certainly anomalous in the genus. The flowers are less strongly arcuate than those of M. maroana, but in characteristics of the calyx and inflorescence the relationship is quite clear. M. ulei is most nearly related to M. neblinensis and differs in having sessile or nearly sessile flowers, and sessile leaflets on a relatively short petiole. Fig. 22. Myllanthus. a-d, M. maroana (Maguire, Wurdack & Keith 41780). a, one leaf, xX 0.25; b, portion of stem with ramiflorous inflorescences and infrutescences, X 0.5; e, flower at anthesis, X 2; d, one mature coceus of fruit, K 2; e-f, M. neblinensis (Maguire, Wurdack § Maguire 42001). e, leaflet, * 0.5; f, inflorescence with immature flowers, < 0.5. APR. 1960 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN 10 (2) :38-57 THE MORPHOLOGY AND RELATIONSHIPS OF DIOMMA, GEN. INC. SED. Wiuuiam L. STERN AND GEORGE K. BrizicKy Samuel James Record Memorial Collection School of Forestry, Yale University New Haven, Connecticut INTRODUCTION In natural systems of plant classification, an attempt is made to arrange plants so that related taxa are near to one another. The bases upon which deci- sions are made concerning relationships among plants are various, and features that are important in some groups are relatively unimportant in other groups. It is now regarded of utmost importance in phylogenetic considerations to select not a single character upon which to found relationships among taxa, but to obtain information from as many pertinent fields of endeavor as possible, and to bring these data to bear upon problems in classification. This synthesizing of available information has been accomplished with more or less success for a number of taxa. Nevertheless, there are still many enigmatic, incompletely investigated groups which have remained in a state of taxonomic limbo affilated at one time with one category and at other times with other categories. Such has been the case with Diomma, genus incertae sedis, noted by Engler from speci- mens collected by Ule in 1910, but actually published by Harms (1931a). It was suggested recently by Dr. Bassett Maguire of The New York Botanical Garden, that a combined morphologic-taxonomie study of Diomma be made in an effort to determine its relationships. In the course of correspondence regarding specimens of Diomma, Dr. Richard S. Cowan of The United States National Herbarium recommended that we also investigate the genus Sohnreyia in this regard. Dr. Cowan remarked, that at least superficially Diomma and Sohnreyva appeared remarkably alike. At the present time descriptions of two species of Diomma appear in the literature, and it will remain to be shown whether these represent distinct taxa and also whether other species perhaps exist. Diomma-» Diomma ulei Engler ex Harms remained the only species until 1952, when Steyermark described a second species, D. fruticosa. Diomma comprises trees of striking appearance for dicotyledons. The unbranched trunk is surmounted by a crown of large pinnately compound leaves and the trees greatly resemble palms in habit. According to Maguire’s field observations, at maturity the plants are crowned by a large compound: paniculate inflorescence. Apparently the plants die after the fruits mature. Harms (1931a) describes Diomma ulei as. possessing large pinnately com- pound leaves. Flowers are white, regular, bisexual, 5-merous, arid borne in short panicles (sic). The free stamens oppose the sepals, and are broadened at the base of the filament. The broad ovary is laterally compressed, 2- or rarely 3- locular, and borne on a broad, cushion-like, nectariferous base. Locules are uniovulate. Ovules are anatropous, and hang from the upper portion of the septum on a short, thick funiculus so that the raphe is dorsal and the micropyle inside (fig. 1A). This condition has been termed apotropy. Styles are 2, very short and stout, each terminated by a capitate stigmatic surface. Samaras are Rott ine 1960 | MORPHOLOGY AND RELATIONSHIPS OF DIOMMA 39 Fic. 1. A, Diomma species A, pistil from female flower to illustrate position of secretory cavities s and the apotropous orientation of the ovules as seen in cleared preparations. B, D. ulei, pistil from female flower to show position of secretory cavities s and nectaries n. C, Diomma species A, pistil from female flower to illustrate shape (ef. A), position of seecre- tory cavities s and presence of nectaries n. D, Spathelia subintegra, pistil from female flower to demonstrate the tricarpellate nature of the ovary, presence of secretory cavities s, and nectaries n. E, Diomma species A, pistil from male flower showing a reduced ovary supported by a gynophore g. F, Sohnreyia excelsa, young fruit showing secretory cavities s and con- Spicuous gynophore. Notice also the beginnings of wings and the peculiar apical protuberance. All drawings diagrammatie. 40 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 2. rarely 3, winged, flattened, broadly ovate, rounded at the base, and attenuated at the tip into a bifid apiculum. Seeds lack endosperm and have a straight embryo. Harms (1931b) added that he did not observe any resin eanals, that the bark and leaves were weakly bitter, and that tufted hairs occurred on the edges of sepals and petals. Steyermark’s (1952) description of D. fruticosa, based on non-flowering material, is essentially the same as that for the genus as given by Harms (1981la), but he mentioned the occurrence of punctae in the leaves. The geographic distribution of Diomma appears to be restricted to the Roraima sediments of the Guayana region of north-central South America. Ule (1914), in describing the vegetation of Roraima, mentioned ‘‘. . . eine merkwurdige neue Gattune der Simarubaceen, Diomma Ulei Engl.’’ Harms (193la), in validating Engler’s description of this species, remarked that the position of Diomma is uncertain. He stated that the apotropous ovules differ from those of genuine Simaroubaceae and from those of the Geraniales as a whole. However, he placed it under Simaroubaceae because Engler had consid- ered it under that family. Nevertheless, later in 1931, Harms delved further into the relationships of Diomma and averred that the apotropous ovules sug- gested an alliance with Sapindales, but that a close relation in this order would be difficult to find. Diomma was placed in the simaroubaceous tribe Picram- noideae next to Alvaradoa by Cronquist (1945). Steyermark (1952) mentioned that, ‘‘The large leaf-scars and long-pinnate leaves of this genus resemble some genera, such as Ailanthus, of the Simarubaceae ... but the punctate leaves of D. fruticosa may show a closer affinity or connection with Rutaceae, to which family it is here assigned.’’ Sohnreyia. The genus Sohnreyia is represented by a single species, S. excelsa, which was described by Krause (1914) from specimens gathered by Ule. Sohnreyra like Diomma, is one of those dicotyledons with a palm-like habit. Krause describes this plant as an erect, palm-like tree with a slender unbranched trunk bearing very large pinnate leaves crowded at the apex. Also, ike Diomma, Sohnreyia iS a Species which blooms and fruits but once at the termination of its existence (Krause 1921; Ducke 1930). Krause (1914) notes that the large, pinnately compound leaves of Sohnreyia are glandular-punctate. Flowers are white, regular, polygamous, 5-merous, and disposed in loose, many-flowered panicles. The free stamens are inserted at the base of a gynophore opposite the sepals. In the female and bisexual flowers, stamens are smaller than in the male flowers. The filaments are provided with a wing-like, bifid appendage in the lower part. The 2-locular ovary is inserted on a thick, subcylindrical gynophore. Ovaries are small, rudimentary, and estigmatose in male flowers, but in female and bisexual flowers the ovaries are larger, somewhat laterally compressed, and in females bear a short style termi- nated by a fairly large, bilobed, discoid stigma. Ovules are pendulous, ana- tropous, and solitary in each locule of the ovary. Krause did not see fully developed fruits. However, in 1922, Ducke described the fruits of Sohnreyia as resembling the samaras of certain Combretaceae, e.g., Terminalia argentea. However, he noted that the fruits of Sohnreyia were 2-seeded, dry, and with 2 membranous confluent wings which are conspicuously reticulate. According to Ducke and Black (1953), Sohnreyia excelsa is scattered in Brazil on uplands 1960] MORPHOLOGY AND RELATIONSHIPS OF DIOMMA 4] from Manaus to the lower part of the Trombetas River north of the Amazon River, and south of the Amazon River from Juruti Velho and Maués to the northern part of the Mato Grosso and the Territory of Guaporé. Krause (1914) placed Sohnreyra in the Rutaceae, in an isolated position, mentioning that it could be related to Dictyoloma, but lacking mature fruits no definite association could be made. Engler (1931) agreed with Krause regarding the definitely rutaceous nature of Sohnreyia and erected a special monotypic subtribe Sohnreyiinae under the tribe Toddalieae of the subfamily Toddalioideae to contain this genus. METHODS AND MATERIALS The investigation undertaken herein depended upon an examination of sev- eral kinds of materials, some by more than one method. Most material was in dry form, but some formalin—acetic-acid—alcohol preserved material was also available for study. Nodal regions and vasculation of petioles in Diomma were examined using a supersaturated solution of phloroglucin in water to demon- strate the presence of lignin. Freehand serial sections were cut directly into the phloroglucin solution, mounted on a slide, and 2 or 3 drops of concentrated HCI applied directly to the section. The appearance of a red-violet color indicated the presence of lignin (vascular tissue). Secretory structures, in leaves of - various specimens examined, were demonstrated by the use of small portions of leaflets cleared by treatment with 3 per cent NaOH at 40°C. Subsequently to washing with water, cleared leaves were stained in safranin, dehydrated, treated with toluene, and mounted on slides in Canada balsam. This same method was employed with pistils and other floral parts. Leaves were also embedded in paraffin, sectioned transversely and paradermally, stained in Heidenhain’s iron- alum haematoxylin and safranin, and mounted on slides in balsam. Wood speci- mens were prepared for microscopic examination in the conventional way. Small blocks were boiled in water to exclude air and soften, after which they were dehydrated and embedded in celloidin. After being cut on a sliding microtome, sections were stained in Heidenhain’s iron-alum haematoxylin and safranin and mounted on slides in balsam. Maceration of wood was accomplished by Jeffrey’s method as outlined by Johansen (1940). Macerated wood was stained in safran- in, dehydrated with tertiary-butyl alcohol, and infiltrated with Canada balsam before being mounted on slides. Studies of floral morphology were based largely on boiled flowers from herbarium specimens. Some preserved flowers were also at hand. Terminology used in the section on xylem anatomy follows that recommended by the Committee on Nomenclature of the International Association of Wood - Anatomists (1957). Terms referring to cell size are taken from Chattaway (1932). Measurements of vessel element and fiber-tracheid length and tangential pore diameter are based on 50 samples each. Only the means and ranges for these measurements are presented because the small number of specimens at our disposal would make meaningless any more detailed treatment of the data (Stern & Greene 1958). Specimens examined: Diomma ulei Engler ex Harms: E. Ule 8646 (Type, G; Isotype, L, K); J. A. Steyermark 58982 (F, NY), 59798 (F). D. fruticosa Steyermark: J. A. Steyer- mark 60820 (Type F sterile), 60327 (‘‘Co-type’’ F fruits); 7. Lasser 1786 (US); B. Maguire & J. J. Wurdack 33808 (NY). Diomma species A: J. A. Steyermark 75699 (NY): 42 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 J. A. Steyermark & J. J. Wurdack 938 (US), 1099 (NY). Diomma species B: B. Maguire, J.J. Wurdack § .C. Maguire 42329 (US, Y), 42315 (US, Y). Sohnreyia excelsa Krause: A. Ducke 8 (Y), 1180 (US, Y), s. nm. (US, MG 15702), s. n. (US, RB 20472). Wood specimens were available of Diomma species B, B. Maguire, J. J. Wurdack & C. Maguire 42315, and of Sohnreyia excelsa, A. Ducke 8. Preserved material of Diomma was available as follows: fruits and petiole portions, Bb. Maguire, J. J. Wurdack & C. Maguire 42329; flowers, leaf and petiole portions, B. Maguire, J. J. Wurdack & C. Maguire 42315; flower buds, J. A. Steyermark & J. J. Wurdack 1099. GENERAL MORPHOLOGY In the course of our investigations several new structural features and ar- rangements of sex in Diomma and Sohnreyia have come to light. Because we probably have had more specimens for study than had Harms, Engler, or Krause, it 1s possible to show certain heretofore unreported characteristics to g2ood advantage. For the sake of the discussions which follow, it is of importance to set down these additions to the knowledge of the genera in question. Diomma. 1. All species of Diomma apparently are polygamous. However, because of the relative paucity of flowering material, the kinds of flowers that occur in the different species are uncertain. In D. ule only female flowers were found; in D. fruticosa and Diomma species B, only male flowers were seen. In Diomma species A, both male and female flowers were observed in the same inflorescence. 2. Sepals, and often petals, are provided with a tiny glandular cavity at their apices. . 3. The filaments of stamens in species of Diomma possess 2 lateral, wing-like expansions (appendages?) in the lower half (fig. 2A—D). These have free, cil- iate, dentiform tips and are usually densely villous on the inner surface. Unfor- tunately, only female flowers of D. ulei were on hand, and the staminal filaments (stamens here are apparently non-functional) here did not possess distinct wing-like portions. However, the bases of the filaments were more or less slightly broadened and ciliate (fig. 2A, B). 4. (varies of female flowers (fig. 1) are subsessile and are situated on a tiny pedestal or rudimentary gynophore. This pedestal bears 5, unequally large, ecushion-like swellings. These are sometimes more or less confluent and appar- ently are nectariferous. A single very short style which terminates in 2 large, discoid to subeapitate stigmatic lobes is present. The stigmata are fleshy and papillose, and may be coherent or actually connate so as to produce a single, deeply bilobed structure. Stamens are somewhat shorter in female than in male flowers. Male flowers possess pistils with reduced ovaries and rudimentary stig- mata. Interestingly, in these flowers, the ovarian pedestal, which is inconspicu- ous in female flowers, forms:a more or less conspicuous, broad, gynophore. : 5. Within the dorsal wall of each carpel there is an elongate secretory cavity, narrow at the apex and more or less bulbous at the base, which is filled with yellowish contents (fig. 1A—C). | There has been some doubt cast on the presence of certain characters in Diomma upon which we can now elaborate. For example, there is no question that the pendulous ovules of Diomma are indeed apotropous, that the raphe is dorsal, and that the micropyle is directed upward (fig. 1A). 1960] MORPHOLOGY AND RELATIONSHIPS OF DIOMMA 43 SRO ae tt tec Wt De oe a ai fiery at Ch A be Fs. Ass Fig. 2. A-D, Diomma spp., stamens to illustrate nature of staminal appendages. A, B, D. ulei from female flower. C, Diomma species A, from female flower. D, Diomma species B, from male flower. E, Sohnreyia excelsa stamen showing structure and orientation of ala as compared with those in Diomma, A—D, and Spathelia, F-I. F-I,Spathelia spp. show- ing variable nature of staminal alae in this genus. F, S. cubensis from male flower. G, S. subintegra from female flower. H, S. simplex from male flower. I, S. glabrescens from male flower. All drawings diagrammatie. 44 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Ule, collector of the type of Diomma, did not mention its palm-like appear- anee. The only report in the literature of this characteristic is in Steyermark’s description of D. fruticosa. However, field notes associated with the following collections also make note of the palm-lke nature of Diomma: D. ulei, J. A. Steyermark 59798; D. fruticosa, T. Lasser 1786, B. Maguire & J. J. Wurdack 33808; Diomma species A, J. A. Steyermark 75699, J. A. Steyermark & J. J. Wurdack 938. Monocarpy, that is, the condition in which death of a plant follows closely after a single flowering and fruiting cycle, has only been noted once for Diomma —in the field notes accompanying B. Maguire & J. J. Wurdack 33808, D. fruticosa. More recently, in correspondence, Maguire has supplied the follow- ing clarification : ‘‘ Apparently the tree does not survive flowering and fruiting, and dies after the fruits have become mature.’’ Further indirect evidence is obtainable in the field notes of 7. Lasser 1786 that the inflorescence is terminal, and in J. A. Steyermark 75699 that the tree is common in the sterile condition. Sohnreyia. 1. Ovules in Sohnreyia are apotropous, the raphe is dorsal, and the micro- pyle is directed upward. 2. The locules of ovaries and young fruits are pubescent inside and possess unicellular trichomes. 3. Sepals are provided with tiny glandular cavities at their apices. 4. Ovaries and young fruits are characterized by the presence of 2 dorsal, elongate, sac-like secretory structures, one in each carpel wall. 5. In Sohnreyia, staminal filaments (fig. 2E) possess wing-like processes at their bases. These processes appear to be fused with the ventra of the filaments for most of their length, their bifid apices being free and ligular and projected ovary-ward. The actual nature of staminal processes here, and the expansions of the filaments in Diomma, cannot be stated with certainty without more de- tailed anatomical studies. NODE AND PETIOLE Although a number of specimens of Diomma were available for study, only one, D. ilei, J. A. Steyermark 58982, proved to be more or less suitable for ana- tomical study. Difficulty is encountered in interpretation of the vascular struc- ture of the stem tip because of the crowded insertion of the leaves and corre- spondinely telescoped internodal regions. However, it can be seen that Diomma has multilacunar nodes with probably 9-15 traces entering the base of the petiole (D. ulei-9, 11; Diomma species A, J. A. Steyermark 75699-1383, 15). Trace num- ber may even vary in a single specimen. The vascular tissue in the petiole was traced from the point of attachment to the stem, the lower pulvinus, to the insertion of the first pair of leaflets, the upper pulvinus. In the first 10 or 12 mm of petiolar tissue, the originally discrete vascular strands fuse (fig. 3B). The resulting vascular cylinder is deeply notched adaxially and corresponds to the adaxial invagination toward the base of the petiole. Distally, the cylinder becomes almost circular in transverse section and is only shallowly notched above. This is true even though the petiole appears quite circular externally in this area. Lignification and hardening of the vascular cylinder seem to proceed from the abaxial to the adaxial regions. It may also be of importance to note 1960 | MORPHOLOGY AND RELATIONSHIPS OF DIOMMA 45 the formation of a sclerenchymatous zone exterior to (part of ?) the phloem and becoming visible 6 or 8 mm distal to the petiolar attachment. This zone first develops in the abaxial region as an are of sclerenchyma, and as one pro- ceeds distally, the are extends until it surrounds the vascular cylinder. No medullary vascular bundles were observed in the pith of the petiole of Diomma. SECRETORY STRUCTURES Secretory structures in the tissues of plants can be divided into two general categories: (1) the secretory cell which is idioblastic in nature and is not con- tiguous with other such cells, and (2) the secretory intercellular space which is surrounded by a continuous layer of cells composing a secretory epithelium. AK OXYYY wane RM i) \ SY Sd sS Bg jee) aS Fic. 3. A, Sohnreyia excelsa, diagrammatic representation of secretory cavity from leaf to show association with vascular tissue and illustrate the general nature of these structures in the taxa under consideration. B, 1-4, Diomma ulei, serial sections of petiole from lower pulvinus at 1 to upper pulvinus at 4, demonstrating the formation of a vascular cylinder from originally distinct vascular strands at 1 (a = vascular tissue, b = selerenehymatous zone). , 46 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Secretory cells retain the ergastic substance (e.g. oil, mucilage, resin), whereas cells in a secretory epithelium release ergastic material, by one means or another, into the intercellular space. A secretory intercellular space which is pocket-like and of more or less definite shape and size is termed a secretory cavity (Stern 1954). Secretory cavities occur in the pistils and leaves of Diomma and Sohn- reyu. Secretory cavities in the pistils of Diomma (fig. 1A-C) and Sohnreyia are essentially similar in form, number and location. Two cavities are present in each pistil, one in the median dorsum of each carpel, close to the outer wall. The secretory cavities are elongate, tapering gradually from the more or less bul- bous base toward the attenuate apex. A yellowish material was observed in these pouches in one specimen. The leaves of Sohnreyva and all species of Diomma are characterized by the presence of secretory cavities in the mesophyll. These cavities vary from 72 to 172 w in diameter and average about 100 p. The secretory epithelium appears to be 1 or sometimes 2 cell layers thick. Cavities nestle within a basket-like net- work of vascular bundles (figs. 3A, 4). In addition to secretory cavities, leaves of all species of Diomma possess eloneate secretory cells distributed amone the chlorenchymatous cells of the palisade mesophyll (fig. 4). These are larger in paradermal section than the palisade cells and average 26 » in diameter, ranging from 14 to 47 p. In length they approxi ate that of the palisade cells. The distribution of secretory structures in the available specimens of the two genera is Summarized in the key below. It should be emphasized that the positions given are apparently not immutable, but may vary to a certain degree. For example, in species described as having secretory cavities located along the margin of the leaflet, an occasional cavity may occur elsewhere in the leaflet. 1. Secretory cells present in leaves and scattered among the chlorenchymatous cells of the palisade layer. Diomma. 2. Secretory cavities confined to the margin of the leaflet. “‘Diomma uler distribution’’: EH. Ule 8646; J. A. Steyermark 75699, 58982, 59798; J. A. Steyer- mark & J. J. Wurdack 938, 1099; B. Maguire, J. J. Wurdack & ad C. Maguire 42315, 42329. j 2. Secretory cavities dispersed throughout the mesophyll of the leaflet. “Diomma fruticosa distribution’’ : J. A. Steyermark 60820; B. Maguire & J. J. Wurdack 33808. 1. Secretory cells absent in leaves. Sohnreyia. XYLEM ANATOMY Whether the concentric marks that appear on the transverse sections of both Diomma and Sohnreyia (fig. 5C) are related to seasonal increments in stem diameter is unknown. It is probably best therefore, to refer to these lines in a general way as concentric growth marks (Chowdhury 1939). Concentric growth marks in Sohnreyia are regularly spaced and obvious because of a pronounced thickening in the walls of the imperforate tracheary elements and the presence of a band of axial parenchyma associated with these thicker-walled fibers. Growth marks in Diomma are irregularly spaced and seem to be erratically pro- duced. Some of these rings are visible because of their delimitation by a narrow band of axial parenchyma (fig. 5A) and/or close tangential aggregations of 1960 | MORPHOLOGY AND RELATIONSHIPS OF DIOMMA 47 ee gl ght 4. - jt we Fig. 4. A, Diomma species B, transection through leaf to illustrate marginal secretory cavities s and secretory cells c in palisade mesophyll, &K 140. B, D. fruticosa, paradermal sece- tion of leaf showing secretory cells c among chlorenchymatous cells of the palisade mesophyll and seattered secretory cavities s, X 180. 48 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vont. 10 pores. However, it is felt that this particular specimen is abnormal or juvenile as there is a fairly wide conspicuous band, 1-3 mm wide, near the center of the stem which is composed of great numbers of closely placed pores and abundant paratracheal parenchyma. This concentric band resembles the early wood of ring-porous species. It is possible, therefore, that were it not for the effect of this seemingly abnormal or juvenile condition on the remainder of the xylem cylinder, the concentric growth marks elsewhere in the stem of Diomma would be of regular occurrence as in Sohnreyia. Both genera exhibit diffuse-porous wood. Imperforate tracheary elements in both genera are fiber-tracheids bearing pits with rounded borders and extended inner apertures. Length varies from very short to short (table 1; Chattaway 1932). In Diomma and Sohnreyia pores (fig. 5A, C) are mostly distributed as radial multiples (2-10 cells), over half appearing in this form. It is interesting to note the occurrence of oc¢ca- sional chains-.of pores in each genus (table 2). Pores are circular in outline and extremely small to moderate-sized in diameter (table 1; Chattaway 1932). Vessel elements are short to very long, and end walls are oblique, but generally not less than 45° (table 1; Chattaway 1932). Intervascular pitting is strictly alternate, although in Diomma there is a local tendency to opposite pitting. Perforation plates are simple, but in Diomma vestigial bars in the form of mucros were occasionally observed projecting from the rim of the plate. Vascular rays are generally heterocellular in Sohnreyia (fig. 5D) the multi- seriate rays exhibiting upper and lower uniseriate ‘‘wines’’ varying from a single marginal cell to 7 cells high. Vascular rays in Diomma (fig. 5B) are essen- tially homocellular and consist mostly of procumbent cells. The sporadic oceur- rence of multiseriate rays with terminal uniseriate ‘‘wines’’ was noted, but cells in these wings were generally procumbent. Occasionally a small ‘‘square’’ cell was observed in the radial view. Uniseriate ‘‘winegs’’ are mostly less than 6 cells in height. Vascular rays in Sohnreyia are regularly triseriate, with some biseriate rays. The uniseriate and quadriseriate conditions are rare. In Diomma the biseriate ray is most abundant with uniseriate rays of fairly common occurrence. Tri- Serlate réiys are very scarce. Axial xylem parenchyma in both genera occurs apo- and paratracheally. In Sohnreyia (fig. 5C) concentric bands 2-3 (4) cells wide are of regular occur- rence. A few strands of parenchyma were also noted in association with vessels or groups of vessels. Concentric bands of axial parenchyma also appear in DPiomma (fig. 5A), but these are mostly 1 cell wide and are irregularly distrib- uted on the transverse section. Incomplete sheaths of axial parenchyma, 1-2 cells wide, are associated with the vessels. : It is apparent from the above description that the xylem anatomy of Sohn- reyia and Diomma is very similar. A few points of difference are however noted in the following features: (1) concentric bands of axial parenchyma are 2-3 cells wide in Sohnreyia and only 1 cell wide in Diomma, (2) vascular rays are usually heterocellular in Sohnreyia and essentially homocellular in Diomma, and (3) vascular rays are regularly triseriate in Sohnreyia and biseriate in Diomma. In considering the significance of these differences thought should be given to the fact that only one wood specimen of each genus was available for analysis, i al € and i and x bi- bd * < ae a at s, X uniseriate banded a illustrating tion illustrating uni- sel element ondition of the banded axial DIOMMA show f ves wide ¢€ al see € Cc OF SO ell D, Tangential section nsection to Tangenti ll b oblique end wa -@ a —4 b] 9” 4 B oe en? 9" tam A 88 Oe Pagel cht ee eeg): get Some eg. i a na ag ge * es * ood. and , Transection to show Ss dp Ay — lam — NM Z eo) _ H m, 4 ea) iow a Z all es) s B, w v # eee se & & $e ae $e AR te I ray r a oe C * & & * & ee % COC we CEC * aseul MORPHOLOG tT. Vv Diomma specie Sohnreyia excelsa, wood. b] and pores in radial multiples. B SSS EOE LTE Se EES eS bd A ma 7p, ‘ Se | ni al TT At Tt Jor See eB Oe S PS ‘Wie. thesia f & ee @ WLS LH SL ca a MS APES Oe ER RS Pe ial. eo ‘ ¥ ry ~ € * = s ~ ~ S322) & a Est * 5. Ais, hy Fig. CS & o's & ea” eee PD parenchyma p, and pores in radial multiples triseriate vascular rays, X 98. biseriate condition o parene 50 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN’ |vou. 10 and it is probable that an abnormality exists in the Diomma specimen and/or it does not exhibit mature characteristics. Table 1. Dimensions of tracheary elements. Diomma Mean Range Fiber-tracheid length 786 uw 405-1200 uw Pore diameter 63 pu 29-104 uw Vessel element length 538 uw 300-825 w Angle of vessel element end wall 54° ‘— tD—-30- Sohnreyia Fiber-tracheid length 901 uw 450-1170 uw Pore diameter 97 pw 40-165 pu Vessel element length 509 wu 360-780 uw Angle of vessel element end wall 54° 80-10° Table 2. Pore distribution in per cent. Solitary Radial multiples Clusters Chains Diomma 36.4 53.0 10.6 few Sohnreyia 34.3 60.0 5.7 few DISCUSSION Two species of Diomma, D. ulei and D. fruticosa, and Sohnreyia excelsa have been validly described in the literature and represent distinct biological entities. In the course of our researches, we were able to examine more specimens assigned to each of these genera than were available to previous workers. Examination of these specimens brought to hght only the single heretofore described species of Sohnreyia. However, besides the two previously described species of Diomma, material of at least two further species was present among our specimens. In this work these have been designated for convenience Diomma species A and Diomma species B. Collections representative of these are listed at the end of the section on methods and materials. Relationships of Diomma. It is the contention of the authors that the characteristics of Diomma defi- nitely indieate its placement among the Rutaceae. The general structure of flowers and fruits is rutaceous in all respects except for the presence of apo- tropous ovules. This latter feature, however, also occurs in the genera Spathelia and Sohnreyia, both of which have been included in Rutaceae by Engler (1931). However, Harms (1931la) considered the presence of apotropous ovules as a characteristic that argued against joining Diomma with any of the families of Geraniales (sensu Engler). | If we were to extend Harms’ thesis on ovule position in Diomma to other groups and exclude certain taxa on the basis of this single characteristic, many unnatural relationships would result. Consider Rhamnales: Rhamnaceae possess epitropous ovules, but Vitaceae and Leeaceae are distinguished by apotropous ovules (Suessenguth 1953). Most Meliaceae are epitropous, but Cedreloideae are apotropous; most Simaroubaceae are epitropous, but Alvaradoideae are apotropous. Within other families, ovule position varies among the genera. This is true for example, in Theaceae (Melchior 1925) and Guttiferae (Engler 1925). Apparently the taxonomic significance of ovule position, as well as of other 1960] MORPHOLOGY AND RELATIONSHIPS OF DIOMMA D1 characteristics, depends upon the taxa in question. In certain groups it may be important only in characterizing subfamilies, in other groups it may be impor- tant at the family level, and it is entirely possible that it may have little or no taxonomic importance whatsoever. It seems to the authors, therefore, too restric- tive to exclude Diomma from Rutaceae on the basis of a single characteristic when so many other features of the genus indicate its rutaceous nature. The occurrence of secretory elements in the leaves of Diomma points to a union with Rutaceae. Blenk (1884) was among the first to report the occurrence of secretory cells in leaves of Rutaceae, namely in Spathelia simpler L., consid- ered in Simaroubaceae at that time. About 20 years later, Schulze (1902) men- tioned secretory cells in leaves of Zanthorylum fagara lL. Neither Solereder (1908) nor Metcalfe and Chalk (1950) mention the occurrence of these cells in leaves of Rutaceae except for Solereder’s citation of Schulze’s work. The secretory structures that are obvious when one examines leaves of Rutaceae are secretory cavities; secretory cells are visible only with the aid of the compound microscope. Engler (1931) states that the secretory cavities that occur in Rutaceae serve to distinguish this family from the closely related Simaroubaceae and Meliaceae. The authors would like to suggest that the occur- rence and distribution of secretory cells and their relationship to secretory cavities within Rutaceae may be of considerable taxonomic importance. Although in our study we have been able to examine relatively few speci- mens, statistically speaking, several observations are perhaps worthy of note: (1) The presence of secretory cells in the leaves of Diomma serves to distinguish this genus from Sohnreyia. In this regard, the occurrence of secretory cells seems to be of generic significance. (2) The distribution of secretory cavities, on the other hand, may be of specific value, at least in Diomma. In the ‘‘ Diom- ma uler distribution,’’ secretory cavities are confined for the most part to the margins of leaflets; in the ‘‘Diomma fruticosa distribution,’’ secretory cavities are dispersed throughout the mesophyll of the leaflet. Xylem anatomy in Diomma resembles that of many rutaceous species. The presence of pore chains and abundant radial pore multiples, in particular, 1s characteristic of a number of Rutaceae. Furthermore, woods of Rutaceae as a whole are relatively consistent in their wood structure with respect to the occur- rence of diffuse porosity, simple perforation plates in the vessels, alternate inter- vascular pitting, axial parenchyma which in many cases is scanty paratracheal with or without terminal or non-terminal bands, and non-septate fibers with simple or narrowly bordered pit-pairs (Heimsch 1942) Pollen morphology of Diomma does not contradict its placement in Rutaceae. A preliminary report of investigations (G. Erdtman, in correspondence) indi- cates that ‘‘The pollen grains in Sohnreyia excelsa, in Diomma sp. and Spathelia subintegra (as well as in S. cubensis) have certain characteristics in common with those of Dictyoloma and in Evodia duwiellia (possibly also with those in Chloroxylon and Chorilaena).’’ The condition of monocarpy, that is, where death of the plant follows ftlower- ing and fruiting, has been reported in Rutaceae, especially in the genera Sohn- reyia (Ducke 1930; Krause 1921) and Spathelia (Marie-Victorin 1948; Fawcett & Rendle 1920; Wilson 1911). As far as the present authors are aware, this condition does not obtain in Simaroubaceae. At this point it might be of impor- tance to note that data on the occurrence of monocarpy among dicotyledons sy MEMOIRS OF THE NEW YORK BOTANICAL GARDEN ’ [vou. 10 are scarce. The condition is also known in certain aborescent species of Lobelia (Krause 1921) and in many shrubby genera of Strobilanthinae-Acanthaceae (Bremekamp 1944). According to the results of an extensive survey of nodal conditions and leaf anatomy currently under way in the laboratory of Dr. Richard A. Howard,. petioles of Rutaceae are characterized by a ring of vascular tissue without asso- ciated medullary bundles. This condition has been reported above for Diomma. Petiolar anatomy in Simaroubaceae also consists of a ring of vascular tissue, but with or without associated medullary bundles. Plants with the palm-like appearance attributed to Diomma are referred to in German botanical literature as Schopfbdume or tufted trees. The occurrence of this manner of growth among dicotyledons appears to have significance only at the species, or at most the genus level, for it is widespread among the species of many families: e.g. Theophrastaceae (Clavija, Theophrasta), Compositae (Verononia spp.), Caricaceae (Carica), Meliaceae, Simaroubaceae, Sapindaceae, and Anacardiaceae (Harms 1912, 1940; Mildbraed 1922). It should not seem strange, therefore, to place the schopfbaumartig Diomma in Rutaceae, for we already know that Sohnreyia and Spathelia are of this form (Krause 1914; Marie-Victorin 1948), as well as the following Rutaceae: Leptothyrsa (Engler 1931), certain species of Cusparia (Engler 1931), and Dictyoloma peruvianum (Macbride 1949). In order to locate the most natural position for Diomma among the infra- familial taxa of Rutaceae, it has been necessary to review more or less closely the characteristics upon which these taxa have been based. Moreover, when the authors reached a tentative hypothesis in this regard, careful examination of the possible near relatives of Diomma was indicated. Sohnreyia; which will be demonstrated as probably the closest ally, has been analyzed in detail and the results are reported above. Spathelia, a somewhat more distant relative, was studied in less detail. The actual results of the observations on Spathelia will be incorporated in the following sections. Comparison of Diomma with Sohnreyia. Diomma and Sohnreyia are similar to each other in the following respects: (1) both exhibit a schopfbaumartig appearance, (2) both are presumably mono- carpous, (3) secretory cavities are present in leaves of each, (4) the basic struc- ture of the flower and fruit is the same in each genus, (5) both possess apo- tropous ovules, (6) pistils in each genus contain 2 similarly placed secretory cavities, (7) each genus is polygamous, (8) xylem anatomy is very similar in these genera, (9) pollen grains are apparently alike, and (10) pinnately com- pound, estipulate leaves occur in each genus. Despite these many resemblances, Diomma and Sohnreyia are distinet and differ in the following characters: (1) secretory cells are present in leaves of Diomma but are lacking in Sohnreyia, — (2) appendages of staminal filaments in Diomma (except D. uler), appear to be lateral wing-like proliferations of the filaments, whereas in Sohnreyia, staminal processes are fused with the ventra of the filaments for most of their length, but the ligulate apex is directed ovary-ward, (3) ovaries in female flowers of Diom- ma are almost sessile (gynophore rudimentary) but in Sohnreyia ovaries are conspicuously stipitate (fig. 1F), (4) insides of the locules of the ovary and fruit are entirely glabrous in Diomma but pubescent in Sohnreyia, (5) the stel- late hairs characteristic of Diomma have not been observed in Sohnreyna. 1960 | MORPHOLOGY AND RELATIONSHIPS OF DIOMMA D3 Comparison of Diomma with Spathelia. The genus Spathelia contains 14 species and is apparently restricted to the West Indies (Marie-Victorin 1948). It constitutes a monogeneric subfamily of Rutaceae, the Spathelioideae (Engler 1931). This genus has a number of char- acteristics In common with Diomma. Plants of Spathelia are characterized, as are those of Diomma, by unbranched trunks with leaves crowded at the apex—Schopfbawme. Monocarpy is a charac- teristic of Spathelia. Leaves are pinnately compound and the leaflets exhibit marginal secretory cavities and scattered secretory cells. Diomma and Spathelia are polygamous genera. Floral parts in Spathelia are basically pentamerous as in Diomma. Sepals and petals in Spathelia both possess an apical glandular cavity. A gynophore is present beneath the ovaries in Spathelia; it is always very tiny in female flowers, but may be more or less conspicuous in male flowers. Pollen grains of Spathelia resemble those of Diomma. Both genera produce winged fruits . The xylem anatomy of Spathelia vermicosa (Yw; Leon s.n.) is essentially the same as that of Diomma and indeed of many Rutaceae (Heimseh 1942). S. vernicosa exhibits diffuse porous wood. The fiber-tracheids are characterized by minute bordered pits and walls that are from thick to very thick. Pores are distributed mainly as radial multiples some of which contain over 12 pores. Solitary pores are of common occurrence and are more aboundant than pore clusters. Some of the latter are fairly extensive, being 20 or more pores in size. Radial pore multiples verge on pore chain configuration in some instances. The perforation plates are exclusively simple, and vessel element end walls are mostly oblique. Intervascular pitting is strictly alternate and the pits are small with round borders. Vascular rays are heterocellular and 1-3 cells wide. Most rays are uniseriate but biseriate rays are common. Triseriate rays are rare. Although a single wide (+ 12 cells) band of axial parenchyma occurs in our specimen, it is believed to be of traumatic occurrence, since (1) certain of the cells are irregular and abnormal in appearance, and (2) it is the only band present on the sections. Nevertheless, its presence probably indicates the poten- tial of band formation in the genus. Scanty vasicentric axial parenchyma is regularly present. Only a few strands, or none at all, are in contact with vessels on a given transverse section. Many species of Spathelia are characterized by alate staminal appendages more or less similar to those of Diomma (fig. 2F—-I). Other species are devoid of these appendages, much as in Diomma ulet. The shape and means of attach- ment to the filament of these appendages differ with the species of Spathelia, the genus being inconstant in this respect. It might be well to recall at this juncture that in species of Diomma exhibiting filament processes, the latter appear as lateral expansions of the filament, and in this Diomma seems to differ from Sohnreyia whose staminal appendages are probably fused with the ventra of the filaments. We note that within Spathelia both the Diomma and the Sohnreyra types of staminal process are present, as well as other types, and even unappendaged filaments. For example, lateral appendages like those in Diomma occur in Spathelia simplex L. (Stearn 162 as S. sorbifolia lL.) ; seale-like, bilobed appendages oceur in S. cubensis P. Wils. (Wright 2192); fairly large, almost truncate appendages are found in S. subintegra Marie-Vict. (Wright 2191) ; and D4 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN. [vou. 10 unappendaged filaments as in Diomma ulei are found in S. glabrescens Planch. (Stearn 1003). Illustrations of these examples occur in figure 2. Locules in Spathelia as in Diomma are uniovulate. Although Bentham and Hooker (1862) and Engler (1931) mention geminate ovules in the locules of Spathelia, Fawcett and Rendle (1920) state: ‘‘. . . ovules one in each cell... pendulous from the inner angle at the apex of each ovary-cell.’’ The authors have also observed only uniovulate locules in ovaries of female flowers of simplex and S. subintegra. Furthermore, it can be demonstrated that the ovules in Spathelia are apotropous as in Diomma. It is worthy of note in this regard, that even in 1858, Agardh presumed that ovules of Spathelia were apotropous from Gaertner’s description of the seeds of S. simplex (1788). Dorsal, elongate secretory cavities Occur in the carpels of Spathelia (fig. 1D) as in those of Diomma. However, they are more conspicuous in Spathelia, especially in the fruiting condition. Differences; of course, do occur between Spathelia and Diomma. The leaves in Spathelia may be many-foliolate—up to 100-jugate; whereas in Diomma, 22 pairs of leafiets appear at present to be maximal. Leaflets in Spathelia are generally small to medium-sized (up to 15 cm long in S. glabrescens and to 20 em long in S. simplex), and only rarely entire. In Diomma, on the other hand, leaflets are generally large and entire. Aestivation of sepals in Spathelia follows two patterns: valvate (e.g. S. glabrescens and S. subintegra) and imbricate (S. cubensis and S. simplex). In Diomma sepalar aestivation is imbricate. Although glandular cavities occur at the tips of perianth parts in Diomma, they are tiny and inconspicuous compared with those of Spathelia. The ovary in Spathelia is regularly 3-carpellate and 3-locularand only Lats 2-carpellate and 2-locular (Fawcett & Rendle 1920). On the other hand, the ovary in Diomma is generally 2-carpellate and 2-locular, 3-carpellate and 3- locular ovaries being of exceptional occurrence (fig. 1). The fruits of Spathelia are narrowly winged; those of Diomma possess broad wings. Seeds of Spathelia contain a fleshy endosperm, whereas Diomma seeds are lacking in endosperm. The embryo in Spathelia has linear-oblong cotyledons, but these are suborbicular to ovate in Diomma. The Position of Diomma in Rutaceae. From the foregoing discussion it is not too presumptuous to state that Diom- ma has a definite and more or less close relationship to Sohnreyia and Spathelia. Sohnreyia excelsa is the only member of the subtribe Sohnreytinae of the tribe Toddalieae, subfamily Toddalioideae. It might seem reasonable to place Diomma in this subtribe with Sohnreyia, and to raise Sohnreytinae to the rank of sub- family, because of the presence of apotropous ovules in both genera. However, the subfamily Spathelioideae, of which Spathelia is the sole member, already exists in Rutaceae, and it is ,characterized and distinguished from all other Rutaceae by the presence of apotropous ovules. Therefore, aside from two differ- ences amone Diomma, Sohnreyia, and Spathelia to be discussed below, the au- thors can see no reason for not including these three genera in Spathelioideae sensu novo. The only objections which might be raised to the above proposal involve the valvate or ‘‘subvalvate’’ aestivation of sepals, and the endospermous seeds of Spathelia. Sepalar aestivation is imbricate in Diomma and Sohnreyia, and the seeds of these genera lack endosperm. 1960] MORPHOLOGY AND RELATIONSHIPS OF DIOMMA Or Or Regarding aestivation of sepals, we can see no insurmountable obstacle to our scheme, for both types have been recorded in Rutaceae. Engler (1931) mentions aestivation of sepals in Rutaceae as mostly imbricate. Eichler (1878), in his analysis of rutaceous flowers, mentions that open, valvate, and imbricate aestiva- tion occur in almost all subfamilies of Rutaceae. For example, sepals are valvate in Calodendron and Melanococca according to Bentham and Hooker (1862) ; valvate in Bauwerella and mostly valvate in Boroniasect. Valvatae Benth. (Engler 1931). It appears then that in Rutaceae, aestivation of sepals is at most a char- acteristic of generic significance and that related genera may differ in this regard. A few original observations on aestivation in sepals of Spathelia indicate that, contrary to reports in the lterature, aestivation in this genus may be valvate (e.g. S. glabrescens Planch. and S. subintegra Marie-Vict.) or imbricate (e.g. S. cubensis P. Wils. and S. simplex L.). Other authors have used the term *‘subvalvate’’ in referring to aestivation of sepals in Spathelia. Such a term is likely to be confusing and certain to be subjective. In Engler’s general statement of the characteristics of Rutaceae (1931), he notes: ‘‘Samen ... mit oder ohne Nahrgewebe.’’ A survey of Engler’s data indicates that within subfamilies of Rutaceae differences in presence or absence of endosperm are found. A few examples may be pertinent: (1) In the sub- family Rutoideae, most members of the tribe NXanthoxryleae possess exendo- spermous seeds, whereas in the tribes Ruteae and Boronieae members possess endospermous seeds; (2) in the subfamily Toddalioideae, members of the tribe Toddalieae may or may not have endosperm. Thus, the presence or absence of endosperm in the seeds of Rutaceae does not seem to be a constant characteristic of a subfamily and is probably merely of generic importance. From the above, the authors conclude that neither the differences in sepalar aestivation nor the presence or absence of endosperm in the three genera under consideration represents a valid criterion against the erection of the subfamily Spathelioideae to contain Diomma, Sohnreyia, and Spathelia. In regard to our proposal of Spathelioideae sensu novo, Erdtman (in correspondence) states, **. . pollen morphology does not speak against your view of referring Sohn- reyia, Diomma, and Spathelia to the Rutaceae, as a special subfamily.’’ Spath- elioideae, as constituted herein, differs from all other subfamilies of Rutaceae in the presence of apotropous ovules, much as the Cedreloideae differ from other subfamilies of Meliaceae. It appears that within this subfamily, Spathelia is more primitive than either Diomma or Sohnreyia. Several conditions in Diomma and Sohnreyia seem to represent reductions or specializations from that which obtains in Spathelia. For example, although Spathelia regularly has tricarpellate ovaries, both Diom- ma and Sohnreyia regularly have bicarpellate ovaries. Tricarpellate ovaries do appear sporadically in the latter genera, which may indicate a reduction from that state. Seeds in Spathelia are endospermous, whereas endosperm is absent in the seeds of Diomma and Sohnreyia. Wings on the fruits of Spathelia are relatively undeveloped, which may indicate a primitive condition. Fruit wings are broad in Diomma and Sohnreyia. There also seems to be a lack of stability in certain features of Spathelia, which again may be interpreted as a state of primitiveness. This is found in the inconstant type of sepalar aestivation, and in the wide variety of form in staminal processes. It may also be true that the well-developed secretory glands, such as are found in carpels of Spathelia, are 56 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN ’ [vou. 10 less advanced than the smaller, reduced glands in the carpels of Diomma and Sohnreyia. Arguments for and against the primitiveness or advancedness of certain characteristics in plants can sometimes be interpreted in two ways. The above, therefore, is only meant to state the authors’ opinions. Actually the occurrence of the schopfbaumartig habit and monocarpy in the three genera under discussion is based on a relatively small body of information. If it is subsequently shown that these two features are in truth constant char- acters of all species of Spathelia, Diomma, and Sohnreyva, it will strengthen the structure of Spathelioideae sensu novo. Further, more detailed studies, on a wide variety of material, are desirable to determine the exact nature of the staminal filament proliferations described for these genera. CONCLUSIONS 1. The genera Diomma, incertae sedis, and Sohnreyia, Rutaceae, are closely related. 2. Diomma is definitely a member of Rutaceae (sensu Eneler 1931) as is shown by investigations of floral morphology, pollen structure, secretory ele- ments, nodal anatomy, petiole vascular structure, and wood anatomy. 3. Diomma and Sohnreyia should be included in the subfamily Spathel- oideae, sensu novo, as they are both closely related to Spathelia. ACKNOWLEDGMENTS For their interest in suggesting the problem and their help throughout the progress of the study, the authors express their gratitude to Dr. Bassett Maguire and Dr. John J. Wurdack of the New York Botanical Garden and Dr. Richard S. Cowan of the U. S. National Herbarium. The following botanists were very kind in arranging for loans of study specimens from their respective institutions and the authors appreciate their efforts: Dr. Theodor Just of the Chicago Nat- ural History Museum, Mr. William T. Stearn of the British Museum, Dr. G. Taylor of the Royal Botanic Gardens, Kew, Dr. H. J. Lam of the Rijksherbar- ium, Leiden, Dr. Charles Baehni of the Conservatoire et Jardin Botaniques, Geneva, and Dr. John R. Reeder of Yale University. For his kindness in exam- ining pollepyspecimens the authors are much indebted to Dr. G. Erdtman of the Palynological Laboratory in Stockholm. Dr. Richard A. Howard of the Arnold Arboretum gave of his time to explain methods for studying petiole sections. Professor I. W. Bailey of Harvard University deserves especial thanks for his critical reading of the manuscript and his suggestions for improvement. Literature Cited Agardh, J. G. 1858. Theoria systematis plantarum. 1—404. Bentham, G. & Hooker, J. D. 1862. Genera plantarum. 1(1): 1-484. Z Blenk, P. 1884. Ueber die durchsichtigen Punkte in den Blaittern. Flora 67: 49-57, 97-112, 136-144, 204-210, 223-225, 275-283, 291-299, 339-349, 355-370, 371-386. Bremekamp, C. E. B. 1944. Materials for a monograph of the Strobilanthinae (Acanthaceae). Verh. Nederl. Akad. Wet. Sect. 2. 41(1): 1-306. Chattaway, M. Margaret. 1932. Proposed standards for numerical values used in describing woods. Trop. Woods 29: 20-28. Chowdhury, K. A. 1939. The formation of growth rings in Indian trees. Indian For. Ree. II. Utilization 2(1): 1-39. Or —~l 1960] MORPHOLOGY AND RELATIONSHIPS OF DIOMMA Committee on Nomenclature, International Association of Wood Anatomists. 1957. Inter- national glossary of terms used in wood anatomy. Trop. Woods 107: 1-36. Cronquist, A. 1945. Additional notes on the Simaroubaceae. Brittonia 5: 469-470. Ducke, A. 1922. Plantes nouvelles ou peu connues de la région amazonienne. II° Partie. Arch. Jard. Bot. Rio de Janeiro 3: 3-281. 1930. Enumeracao das plantas amazonicas cultivadas no Jardim Botanico e introduzidas pelo chefe de seccao Adolpho Ducke, de 1920 a 1928. Arch. Jard. Bot. Rio de Janeiro 5: 77-98. Ducke, A. & Black, G. A. 1953. Phytogeographical notes on the Brazilian Amazon. Anais— Acad. Brasil. Ci. 25: 1-46. Eichler, A. W. 1878. Bliithendiagramme 2: 1-575. Engler, A. 1925. Guttiferae. In: A. Engler & K. Prantl, Die Natiirlichen Pflanzenfamilien. ed. 2. 21: 154-237. 1931. Rutaceae. Jn: A. Engler & K. Prantl, Die natiirlichen Pflanzenfamilien. ‘ed. 2. 19a: 187-359. Fawcett, W. & Rendle, A. B. 1920. Flora of Jamaica. 4: 1-369. Gaertner, J. 1788. De fructibus et seminibus plantarum 1: 1-284. Harms, H. 1912. Uber die Sapindacee Talisia princeps Oliv. Verh. Bot. Ver. Brandenburg. 53: (36)—-(40). 193la. Gattung zweifelhafter Stellung, vielleicht den Simarubaceen anzu- schliessen. Diomma Engler nach Ule. In: A. Engler & K. Prantl, Die nattrlichen Pflanzenfamilien. ed. 2. 19a: 460. 1931b. Uber die Gattung Diomma Engler. Notizbl. Bot. Gart. Mus. Berlin 11: 282-284. . 1940. Meliaceae. In: A. Engler & K. Prantl, Die natiirlichen Pflanzenfamilien. ed. 2. 19b1: 1-172. Heimsch, C. 1942. Comparative anatomy of the secondary xylem in the “Gruinales” and “Terebinthales” of Wettstein with reference to taxonomic grouping. Lilloa 8: 83-198. Johansen, D. A.» 1940. Plant microtechnique. 1-523. Krause, K. 1914. Rutaceae: Jn: Pilger, Plantae Uleanae novae vel minus cognitae. Notizbl. Bot. Gart. Mus. Berlin 6: 143-149. 1921. Uber einen hapaxanthen Baum. Mitt. Deutsch. Dendr. Ges. 31: 204-206. Macbride, J. F. 1949. Flora of Peru. Publ. Field Mus. Bot. 13: (32): 511-777. Marie-Victorin, Frére. 1948. Nouvelles études taxonomique sur la flore de Cuba. III. Le genre Spathelia (rutacées), avee description de cing espéces et d’une variété nouvelles. Contr. Inst. Bot. Univ. Montréal 63: 14-48, 76. Melchior, H. 1925. Theaceae. In: A. Engler & K. Prantl, Die natiirlichen Pflanzenfamilien. ed. 2. 21: 109-154. Metcalfe, C. R. & Chalk, L. 1950. Anatomy of the dicotyledons. 1—-1500. Mildbraed, J. 1922. Botanik. Wiss. Ergebn. 2‘** Deutsch. Zentral-Afrika-Exped. 1910-1911. 2: 1-718. Schulze, H. 1902. Beitrage zur Blattanatomie der Rutaceen. Beih. Bot. Centr. 12: 55-98, Solereder, H. 1908. Systematic anatomy of the dicotyledons. 1-1182. (transl. L. A. Boodle & F. E. Fritsch). Stern, W. L. 1954. A suggested classification for intercellular spaces. Bull. Torrey Club ( 81: 234-235. ; Stern, W. L. & Greene, S. 1958. Some aspects of variation in wood. Trop. Woods 108: 65-71. Steyermark, J. A. 1952. Contributions to the flora of Venezuela. Botanical exploration in Venezuela—Il. Fieldiana Bot. 28: 243-447, Suessenguth, K. 1953. Rhamnaceae, Vitaceae, Leeaceae. In: A. Engler & K. Prantl, Die natirlichen Pflanzenfamilien. ed. 2. 20d: 7-333, 372-398. Ule, E. 1914. Die Vegetation des Roraima. Bot. Jahrb. 52(Beibl. 115) :42-53. Wilson, P. 1911. Notes on Rutaceae—VI. Species of Spathelia. Torreya 11: 262-264. APR. 1960 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN 10 (2) :58—64 TAXONOMIC RELATIONSHIPS OF DIOMMA ENGLER EX HARMS RICHARD S. COWAN AND GEORGE K. BRIZICKY Department of Botany, U. S. National Museum, Smithsonian Institution, Washington, D. C. and Yale University School of Forestry, New Haven, Connecticut INTRODUCTION The formal taxonomic conclusions reached in this study can hardly be pre- sented without acknowledging the great debt owed Stern and Brizicky for the very lucid treatment of the anatomy and morphology of this group and its rela- tives reported in a preceding paper of this serial.* Much of the history as well as the morphology and anatomy of the groups involved is quite adequately set forth in their contribution and will not be repeated here. However, since three former genera are herein reduced to subgenera of a single genus, the reasons for such a modification should be summarized. The key to the subgenera contrasts the only known differences of any magni- tude ; it is at once apparent that these are few and in most instances more or less continuous, rather than discontinuous as one may justifiably expect differences of generic stature to be. The many similarities among the three taxa are amply sum- arized by Stern and Brizicky and it appears obvious that the similarities far outweigh the dissimilarities. The decision that incipient rather than completely former genera are herein reduced to subgenera of a single genus, the reasons for such a modification should be summarized. The question may be raised that, since this is obviously a monophyletic group, we must account for the present geographic distribution of its parts. If we may place some reliance on the maps presented by Schuchert (1935) showing the configurations of the land masses in and adjoining the Antillean—Carribean region, more or less continuous land masses have been available for migration (via Central America) between Cuba and the Bahamas at one extreme and northern South America on the other for long periods, beginning in the Late Upper Cretaceous and extending well up into the Eocene. Simpson (1940) considers the land connection between North and South America to be a much more recent event—perhaps as late as the Pliocene—but this conclusion does not invalidate the supposition that migration was by a land route between Cuba and northern South America, at least in part. Indeed, the fact that the majority of Cuban species have their closest affinities in South America (Alain 1958) clearly indi- cates rather efficient intercommunication between the two extremes. In addition to migration over land routes, the powerful winds of Caribbean hurricanes must have played an important role in the dispersal of plants in the West Indian— northern South American region. The winged fruits of all species of Spathelia, for example, are susceptible to wind-dispersal, and the present distribution of the genus may be the result of the operation of this agent alone. Whatever mecha- nism was responsible, it is certainly less novel than the direction of the migration, which is unusual but characteristic of Simpson’s ‘‘filter land-bridges’’ (1940), through which migration always occurs in both directions. *10: 38-57. — 58 — 1960 | TAXONOMIC RELATIONSHIPS OF DIOMMA a9 Morphologically, there is good reason to consider subgenus Spathelia the least advanced of the three subgenera (see Stern and Brizicky), and since it is pres- ently restricted to the Bahamas, Cuba, and Jamaica, we may consider this area as the original point from which the genus evolved and migrated southwardly, perhaps by a land-route. Such a view would be strengthened by the presence of some representative of the group in the intervening region (but interconnecting taxa are unnecessary if the dispersal is assumed to be via hurricane winds). A single species from Mexico was referred to the genus by Sessé and Mocino, but Wilson (1911) considers it ‘‘a very doubtful species.’’ Unfortunately, the original publication by these authors is so poor that it is difficult to determine the exact affinities of the plant depicted. TAXONOMY Spathelia Linnaeus, Sp. Pl. ed. 2. 386. 1763. (nom. cons.) (emend. et ampl.) Spathe P. Browne, Hist. Jam. 187. 1756. (nom. rej.) Sohnreyia Krause, Notizbl. Bot. Gart. Mus. Berlin 6: 147-149. 1914. Diomma Engler ex Harms, Nat. Pflanzenfam. ed. 2. 19a: 460. 1931. Monocarpic trees with palm-like habit, the alternate, pinnate leaves congested at the apex of the unbranched trunk. Leaflets opposite to alternate, 10—100- jugate, the margins crenate to entire, glandular punctae restricted to the margins or generally distributed through the blade. Inflorescences terminal, paniculate, to 5 m long, the flowers polygamous. Sepals 5, free to slightly united basally, valvate to imbricate, the apex with a small to large glandular cavity; petals 5, free, imbricate, the apex with a small to large glandular cavity, rarely the latter not obvious. Stamens 5, the filaments exalate to conspicuously alate, the wing shortly free at the apex or completely adnate with the filament, bidentate apically or entire. Gynoecium subsessile to stipitate; stigma 2—3-lobed, sessile or sub- sessile; ovary 2—3-celled, laterally compressed, the carpels uniovulate, the ovule pendulous, apotropous. Fruit 2—3-winged, indehiscent, the wings narrower to broader than the seed-bearing portion; seed endospermous or not. Type species: S. simplex Linnaeus, loc. cit. Key to the Subgenera 1. Ovary typically 3-locular, infrequently 2-locular; fruit elliptic to oval, 3 (—2)-winged, each of the wings usually narrower than the seed-bearing portion; each locule with a large, dorsal secretory cavity; seeds with a fleshy endosperm. Leaflets rarely entire, up to 100-jugate. Plants of the West Indies. Subg. 1. Spathelia. 1. Ovary typically 2-locular, rarely 3-locular; fruit ovate to oblate, 2 (—3)-winged, each of the wings usually much broader than the seedbearing portion; loecules without a large, dorsal secretory cavity; seeds without endosperm. Leaflets entire, to 27-jugate. Plants of northern South Ameriea. 2. 2. Stigma separated from the ovary by a short style, the lobes capitate or subcapitate; gynophore absent or rudimentary; ovary and fruit glabrous within. Secretary cells (microscopic) present in the leaflet tissue. Known only from Guayana. Subg. 2. Diomma. 2. Stigma sessile, the lobes plane; gynophore conspicuous; ovary and fruit densely pubescent on the inner surfaces of the locules. Secretory cells absent in the leaflet mesophyll. Known only from northern Amazonia. Subg. 3. Sohnreyia. Spathelia Subg. 1. Spathelia. The fourteen species included in this subgenus are restricted to the Bahamas, Cuba, and Jamaica; the eleven that are endemic to Cuba are distinguished in a [vou. 10 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN 60 1960] TAXONOMIC RELATIONSHIPS OF DIOMMA 61 partial key and by descriptions presented by Leén & Alain (1951). One further, but doubtful, species from Mexico has been proposed by Sessé and Mocino. Spathelia Sube. 2. Diomma (Engler ex Harms) Cowan & Brizicky, stat. nov. Diomma Engler ex Harms, Nat. Pflanzenfam. ed. 2. 19a: 460. 1931. Type Species: Spathelia (subg. Diomma) ulei (Engler ex Harms) Cowan & Brizicky, comb. noy. (based on Diomma ulei Engler ex Harms, 1.c.) Key to the Species of Subg. Diomma 1. Leaflets with glandular punctae scattered, not limited to the margins. Fruits not over 2.5 em wide, deeply cordate basally, retuse apically, each half about semi- cireular. 1. S. fruticosa. 1. Leaflets with glandular punctae restricted to the margins. Fruits larger, differently shaped. 2 bo Filaments not aliform basally or if slightly expanded, not aliform and not bidentate at the apex. Pedicels +-5.5 mm long. Ovary broadest basally; fruits broadly ovate, attenuate apically, the seed-bearing portion narrowly lanceolate. 2. S. ulei. 2. Filaments winged basally. Pedicels 1.5-2.5 mm long. Ovary broadest at about the middle; fruits oblate or obovate-oblate, not attenuate apically, the seed-bearing portion broadly ovate. De 3. Median. leaflets about 10-12 em long, oblong, acute, the rachis ca. 30-45 em long. Fruits glabrous, oblate, slightly apiculate apically, 4 em broad. 3. S. chimantaensis. 3. Median leaflets about 14 em long, oblong-lanceolate, acuminate, the rachis 47-57 em long. Fruits puberulous, obovate-oblate, truncate apically, 3 em broad. 4. S. neblinaensis. 1. Spathelia fruticosa (Steyermark) Cowan & Brizicky, comb. nov. Fig. 1, a. Diomma fruticosa Steyermark, Fieldiana Bot. 28: 272. 1952. Tree 1—5 m tall, the tip of the stem densely puberulous minutely. Leaves 25-30-foliolate, the leaflets with scattered glandular punctae, the petioles terete except at the canaliculate base, 10-18.5 em long, densely puberulous but glabres- cent, the rachis 42-54 em long, densely puberulous, glabrescent, the leaflets op- posite to alternate, the petiolules 1—-2.5 mm long, puberulous but glabrescent, the blades glabrous at maturity, those nearest the leaf base smaller, ca. 9-10 em long, 3.0—-4 em wide, oblong-ovate, the blades of the upper leaflets larger, slightly to strongly arcuate, oblong-lanceolate or oblong, 9-15 cm long, 3—5 cm wide, the base of all the leaflets inequilateral, the upper side rounded-obtuse, the lower side subobtuse (or rarely acute) to rounded-obtuse, the apex abruptly acute to acuminate with the tip blunt, the venation plane on the upper leaflet surface or the costa slightly salient, the costa and the primary vein-branches salient beneath. Inflorescences 35-63 em long, the axis densely puberulous, the bracts about 1 mm long, puberulous, the pedicels 1-2 mm long, puberulous. Sepals 1.5-2 mm long, 0.8-1.2 mm wide, oval to narrowly elliptic, emarginate apically, glabrous ex- cepting the ciliolate margin; petals 3.5-4 mm long, 2 mm wide, oval, glabrous. Fig. 1. a, Spathelia fruticosa, fruit, X 1 (drawn from Steyermark 60327). b-d, 8S. ulei. b, fruit, X 1; ¢, filament, & 7.5; d, ovary, X 10 (drawn from Ule 8646). e—f, S. neblinaensis. e, leaflet, X 0.5; f, immature (?) fruit, KX 1 (drawn from Maguire, Wurdack §& Maguire 42329). gi, S. chimantaensis. g, fruit, X 1; h, filament, X 7.5; i, leaflet, X 0.5 (drawn from Steyermark 75699). j-m, S. excelsa. j, fruit, X 1; k, lateral view of filament, X 7.5; 1, view of outer face of filament, K 7.5; m, gynoecium, X 7.5 (j drawn from Ducke 20472, k—-m drawn from Ducke 15702). 62 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Filaments (in staminate flowers) 25 mm long, ligulate with a bidentate, alate basal half, the wing 1 mm wide, villosulose on its inner surface. Gynoecium (in staminate flowers) small, rudimentary, resting on and partly immersed in a short, subglobose to broadly cylindric, puberulous gynophore; pistillate flowers not seen. Fruits glabrous, oblate, retuse and minutely apiculate at the apex and deeply cordate basally, 13-17 mm long, 15-25 mm wide. VENEZUELA: Bolivar: Kavanayén, alt. 1200 m, 28 Mar 1946, Lasser 1786 (US); savanna between Kavanayén and base of Ptari-tepui, alt. 1200-1300 m, Maguire and Wurdack 383808 (NY) and 18 Nov 1944, Steyermark 60327 (F); between La Laja and Santa Teresita de Kavanayén, alt. 1220 m, 30 Nov 1944, Steyermark 60820 (F, holotype of Diomma fruticosa Steyermark). Although S. fruticosa was described from fruiting material only, it stands apart quite distinctly from the remaining species on floral characteristics as well. It differs in the distribution of the glandular punctae in the leaflets, in its shorter pedicels, and in the form of the fruit. It is surely most nearly related to S. ulei. 2. Spathelia ulei (Engler ex Harms) Cowan & Brizicky, comb. nov. Fig. 1, b-d. Diomma ulei Engler ex Harms, Nat. Pflanzenfam. ed. 2. 19a: 460. 1931. Tree 6-10 m tall, the tip of the stem puberulous and minutely stellate- pubescent but glabrescent. Leaves 20—28-foliolate, the leaflets with glandular punctae restricted to the margins, the petioles somewhat canaliculate basally but otherwise terete, 17 em long, puberulous, the rachis 45-60 em long, terete, ‘puberulous, the leaflets subopposite to alternate, glabrous, or puberulous on the upper surface of the costa, the petiolules 4-6 mm long, puberulous, the blades of the lower leaflets smaller, ca. 5-6.5 mm long, 2.5-3 em wide, oval to oval-oblong, the blades of the upper leaflets larger and oblong or lanceolate-oblong, 14-17 cm long, 3.5-4 em wide, the base of all the leaflets slightly to very markedly in- equilateral, the lower side acute and the upper rounded-obtuse, the apex abruptly acute to acuminate with the tip blunt, the venation plane to impressed on the upper leaflet surfaces, plane to salient beneath. Inflorescence 10-15 em long, the axis puberulous, the bracts minute, the pedicels 4-5.5 mm long (10-12 mm in the original description), puberulous. Sepals oval, obtuse, entire to emarginate, 3 mm long, 2 mm wide, glabrous except ciliolate; petals elliptic, 5-6 mm long, 4 mm wide, glabrous to sparingly ciliolate. Filaments (in pistillate flowers) 3.5-4+ mm long, hgwiate, linear or subulate or with the basal half somewhat expanded laterally, sparingly villose basally on the inner surface. Gynoecium glabrous, the stigma bifid, each half capitate, the style about 0.3-0.5 mm long, columnar, the ovary oblong-ovate to broadly ovate, ca. 2.5-4+ mm long, 2-3 mm wide, the base broad and with 4 (5?) bulbous sparsely pubescent nectaries. Fruit broadly ovate, acuminate, 2.5-3 em long, 2-38 em broad. VENEZUELA: Bolivar: Mt. Roraima, between Rondon Camp and base of escarpment, alt. 2040-2255 m, 30 Sep 1944, Steyermark 58982 (F, NY); Ptari-tepui, alt. 1800-1900 m, 2 Nov 1944, Steyermark 59798 (F); base of Mt. Roraima, Ule S646 (G, K, type collection of Diomma ulei Engler ex Harms). - Like most of the other species of this alliance in Guayana, S. ulei is poorly known, but it is perhaps most nearly related to S. fruticosa. The latter differs in the shape of its fruit, in the length of its pedicels, and in the distribution of the glandular punctae in the leaflets. In the fragment envelope of the Geneva specimen of the type collection there is a young fruit which is quite different in form from those described by Harms 1960 | TAXONOMIC RELATIONSHIPS OF DIOMMA 63 and from the one complete, mature one in that envelope. The atypical fruit is probably from another plant and in fact is similar to the fruits of S. fruticosa. 3. Spathelia chimantaensis Cowan and Brizicky, sp. nov. Fig. 1, g—-1. Arbor 5-10 m alta, haud ramificata, foliis trunci ad summum congestis, apice juvenili dense puberulo, glabrescenti. Petiol 10-13 em longi, puberuli sed olabrescentes, basim versus canaliculati sed apicem versus plani, foliorum lamina ea. elliptica, rhachi 30-45 em longa. Foliola ca. 25-40, alterna ad subopposita, coriacea, petiolulis 3-4 mm longis; laminae leviter arcuatae, mediales 12 em longae, 3.5-4 em latae, oblongae vel oblongo-ellipticae, ad basim inaequilaterales, latere superiore rotundato, latere inferiore acuto, apicaliter obtuse acutae, supra glabrae, infra leviter ad costam puberulae. Inflorescentiae terminales, pani- culatae, ad 1 m longae, axibus puberulis, bracteis persistentibus, foliaceis, stipitatis, 9-45 mm longis, bracteolis minutis, pedicellis 1.5-2.5 mm _ longis, puberulis et minutissime stellato-pubescentibus. Sepala 1.5-1.7 mm longa, 1-1.2 mm lata, ovalia, minute ciliolata; petala 5.5-6 mm longa, 2.6 mm lata, concava, elliptico-obovata, ad basim cuneata, glabra. Staminorum fllamenta (in floribus masculinis) 4-4.7 mm longis, parte basali alata, alis ca. 2 mm longis, ad apicem bidentatis, ciliolatis, plus minusve villosulis ; ovarium in floribus masculis parvum, rudimentarium gynophorium breve, crassum, subeylindricum vel subglobosum, 1-1.5 mm diametro, dense puberulum insidens, in floribus femineis; nondum ex- pansis) late ovoideum, lateraliter compressum, glabrum, 1.8—2.2 mm longum et 1.2-1.8 mm latum in basi extrema brevi paulo attenuata nectaria 4 (5?), parce puberula gerens; stigma in floribus femineis majusculum, bilobum, lobis capitatis ; stylus ca. 0.3-0.4 mm longus. Fructus oblatus, glaber, 2.5—-3 em longus, 44.3 em latus, ad pedicellum valde cordatus, ad apicem acutus, corpore late ovato. alis coriacels. VENEZUELA: Bolivar: Chimantéa Massif, Torono-tepui; ‘‘tree 5-6 m, occasional. Calyx brick with pale green; petals white; filaments greenish white; anther golden; ovary dull madder with pale green stellate hairs. South-facing forested slopes above valley of South Cafio, on summit,’’ alt. 1955-2090 m, 23 Feb 1955, Julian A. Steyermark & John J. Wurdack 1099 (holotype, NY); same data and collectors 938 (paratypes, NY, US); Chimanté Massif, Apacara-tepui, 19 June 1953, Steyermark 75699 (paratypes, F, NY). The differences that separate this species from S. neblinaensis are not striking, with the exception of the fruiting ones: the fruits are quite dissimilar in size, shape, and vestiture. The two species also differ in size and shape of leaflets. 4. Spathelia neblinaensis Cowan & Brizicky, sp. nov. Fig. 1, e-t. A 8S. chimantaensi rhachi longiore (47-57 em), foliolis medialibus onions et angustioribus (14 em longis, 3-3.3 em latis), oblongo-lanceolatis, ad apicem arcuato-acuminatis, bracteis primariis caducis, petalis latioribus (4 mm), fructu obovato-oblato, 2 em longo, 3 mm lato, dense puberulo, apicaliter truncato differt. Flores femineos non vidimus. VENEZUELA: Amazonas: Cerro de la Neblina, Rio Yatua; ‘‘Tree 7 m. Fruit green. Occasional in Bonnetia serub forest at northwest head of Cafion Grande, alt. 2000 m, summit,’’ 8-9 Dee 1957, Bassett Maguire, John J. Wurdack & Celia K. Maguire 42329 (holotype, US No. 2267446—2267447); same data and collectors 42315 (paratypes, NY, US). In flower the two new species are not easily separable; only the larger, some- what differently shaped leaflets and the longer leaf rachises are reliable char- acters at this stage. Fruiting specimens, however, are easily distinguished by the differences in size, shape, and vestiture of the fruits. 64 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Spathelia Subg. 3. Sohnreyia (Krause) Cowan & Brizicky, stat. nov. Sohnreyia Krause, Notizbl. Bot. Gart. Mus. Berlin 6: 147-149. 1914. Only a single species has been described thus far in this subgenus; since its original description, several additional collections have been made and these furnish the primary basis for the following characterizations. 1. Spathelia excelsa (Krause) Cowan & Brizicky, comb. nov. Fig. 1, j-m. Sohnreyia excelsa Krause, Notizbl. Bot. Gart. Mus. Berlin 6: 148. 1914. Tree 10-20 m tall, the stem tip minutely puberulous. Leaves 30—54-foliolate, the leaflets with scattered glandular punctae, the petioles to 14 em long, canali- culate, deeply so at the base, minute puberulous, the rachis to about 2.5 m long, puberulous, the leaflets subopposite to alternate, the petiolules 2-6 mm long, minutely puberulous, the blades of the median leaflets 23-30 em long, 3.5—5.5 em wide, narrowly oblong to lanceolate-oblong, slightly arcuate, the base strongly inequilateral with both sides rotund or the upper side subobtuse, the apex acuminate with the tip blunt, the costa and 20-30 pairs of primary vein-branches plane on the upper surface, salient beneath. Inflorescence to 3 m long, the axis densely puberulous, the primary bracts foliaceous, narrowly elliptic, ca. 20 mm long, 5 mm wide, the secondary bracts narrowly triangular, about 1 mm long, the bracteoles minute, the pedicels 1-3 mm long, puberulous. Sepals rotund to broadly ovate, 1-2 mm long and wide, sparingly puberulous externally, ciliolate minutely ; petals elliptic, obtuse, glabrous, 4-6 mm long, 2-3 mm wide. Filaments (in pistillate flowers) 2-4 mm long, the basal 1-2 mm alate, the wing bidentate and densely villose, shortly free at the apex from the subulate, apical portion of the filament. Ovary oval, 1.5-3 mm long and wide, glabrous, the stigma sessile, the gynophore 1-2 mm long, densely puberulous. Fruit very broadly oblate, cordate basally, 2.5-3.5 em long, 4.5-6 em broad, glabrous, the seed-bearing portion ca. 8 mm long, 5 mm broad. BRAZIL: Amazonas: Mandaos, Estrada do Paredao, 13 Feb 1943, Ducke 1180 (NY, US); Oriximina, bas Trombetas, Parad, Dee 1915, Ducke 15702 (US); Juruty Velho, Para, 28 May 1927, Ducke 20472 (US). The conspicuous gynophore, the sessile stigma, and the shape of the fruit separate this species from its relatives in the other subgenera. In the shape of the fruit and the distribution of the glandular punctae in the leaflets S. excelsa is most smmilar to S. fruticosa. Literature Cited In addition to the references given in the body of this paper, the following were also consulted: Alain, Hermano (E. E. Liogier). La flora de Cuba: sus principales caracteristicas, su origen probable. Revista Soc. Cub. Bot. 15: 36-59, 84-96. 1958. Harms, H. Ueber die gattung Diomma Engler. Notizbl. Bot. Gart. Mus. Berlin 11: 282-284. 1931. Leon, Hermano (J..S. Sauget) & Alain, Hermano (EB. E. Liogier). Flora de Cuba 2: 389-392. 1951. : : Marie-Victorin, frére. Nouvelles études taxonomiques sur la flore de Cuba. III. Le genre Spathelia (Rutacées), avee description de cing espéces et d’une variété nouvelles. Contr. Inst. Bot. Univ. Montréal 63: 14-48, 76. 1948. Schuchert, C. Historical geology of the Antillean-Caribbean region. i—xxvi, 1-811. 1935. Simpson, G. G. Mammals and land bridges. Jour. Wash. Acad. Sci. 30: 137-163. 1940. Wilson, P. Notes on Rutaceae—VI. Species of Spathelia. Torreya 11: 262-264. 1911. APR. 1960 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN 10 (2) :65-117 ANATOMY OF GUAYANA XYRIDACEAE: ABOLBODA, ORECTANTHE, AND ACHLYPHILA SHERWIN CARLQUIST Claremont Graduate School, Rancho Santa Ana Botanie Garden, Claremont, California The discovery of exceptionally interesting new entities referable to Xydridaceae in the Guayana Highland by Maguire and his colleagues (Maguire, Wurdack et al. 1958, 1960) has led to a reappraisal of the limits of the family and its relationships within the monocotyledons. On another level, there are problems of generic and specific differences and relationships and their evolu- tionary implications. Anatomical studies are important aids in solving these problems. Dr. Maguire, who has organized studies on these taxa, has recognized the helpfulness of anatomical data and has provided me with an excellent col- lection of suitable material. I should like to express sincerest appreciation to him for his gratifying interest in the studies recounted below, his generous sharing of material with me, and his invitation of this contribution. Of par- ticular significance is the fact that the writer reached the conclusions set for- ward at the end of this paper on the basis of anatomical material alone. Dr. Maguire and I exchanged manuscripts when our studies were completed so that we could take advantage of each other’s findings in a final presentation, but the great similarity in our conclusions, based on different methods of study, derives from the fact that anatomical features show the same patterns of likeness and diversification with reference to the taxonomic system as do features of gross morphology. I acknowledge with pleasure Dr Maguire’s interest in my in- dependent development of such facts, and his use of these when they became available to him. If there had been any major divergences of opinion, we should have exposed these and attempted to resolve them, but in fact, no such dis- agreements in interpretation of materials did occur. MATERIALS AND METHODS Both liquid-preserved and herbarium materials were given to the writer. Herbarium material was treated with warm 2.5 per cent aqueous NaOH to expand it to its natural proportions, washed, and stored in 50 per cent ethyl aleohol. Both liquid-preserved and treated herbarium material were embedded, sectioned, and stained according to the techniques described earlier by the writer (1958). Pollen preparation techniques are described below in connection with that topie. The specimens studied are listed below. All specimens are represented by a mounted collection in the New York Botanical Garden Herbarium. Taxa recognized correspond with those of Maguire, Wurdack, et al. (1958, 1960). Liquid-preserved, assorted parts: Abolboda acaulis Maguire, Maguire & Politi 27700, Maguire 32820, Maguire 33730, Maguire et al. 40669; A. acicularis Idrobo & Smith var. acicularis, Maguire et al. 41439; A. americana (Aublet) Lanjouw, Magwire et al. 41537; A. ciliata Maguire & Wurdack, Maguire et al. 42420; A. linearifolia Maguire, Maguire et al. 41680; A. macrostachya Spruce ex Malme var. angustior Maguire, Maguire et al. 41541; A. macrostachya Spruce ex Malme var. macrostachya, Maguire et al. 41446; A. macrostachya Spruce ex = Gace 66 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [von. 10 Malme var. robustior Steyermark, Magwire et al. 28176; A. sprucei Malme, Maguire et al. 41721; Achlyphila disticha Maguare & Wurdack, Magwire et al. 42402; Orectanthe sceptrum (Oliver)Magnire subsp. occidentalis Maguire, Magwre et al. 40567. Liquid-preserved, pollen only: Abolboda bella Maguire, Maguire et al. 34514; A. ebracteata Maguire, Maguire et al. 30467; A. grandis Grisebach var. quayanensis Maguire, Maguire et al. 27561; A. macrostachya Spruce ex Malme var. angustior Maguire, Maguire et al. 36342; A. macrostachya Spruce ex Malme var. robustior Steyermark, Maguire et al. 28176; Orectanthe ptaritepwiana (Steyermark) Maguire, Maguwire et al. 32823, Steyermark & Wurdack 523, Steyer- mark & Wurdack 1221; O. sceptrum (Oliver) Maguire, Maguire et al. 40567, Maguire et al. 33180. Dried materials, assorted portions: Abolboda macrostachya Spruce ex Malme var. macrostachya, Maguire et al. 39957; A. macrostachya Spruce ex Malme var. robustior Steyermark, Maguire et al. 28176; Orectanthe ptaritepuiana (Steyer- mark) Maguire, Maguire 32766, Maguire 32823, Steyermark & Wurdack 908; O. sceptrum (Oliver) Maguire, Maguire et al. 33342, Maguire et al. 40567; O. sceptrum (Oliver) Macuire subsp. occidentalis Maguire, Maguire & Politi 28451. ANATOMICAL DESCRIPTIONS ROOT Solereder and Meyer (1929) summarize the work of Nilsson, Poulsen, and Malme, as well as their own, on root anatomy of Xyridaceae. Although their data apply chiefly to Yyris, with only a few references to Abolboda, they pro- vide a framework for the expression of data. Almost every character they cite is represented by a number of Xyris species, and detailed mention of these, other than acknowledgment of occurrence of those characters in Xyris, is un- necessary here. Certain characteristics, on the basis of the present study, appear to be of greater value than others for systematic purposes. Characters which appear relatively reliable in this respect include: (1) Presence of a continuous pericycle, as opposed to its interruption by vessels. (2) Shape and thickening of endodermis cells. (3) Number of vessels present in the vascular core (only insofar as the number is relatively large or small; a certain variation in the number of xylem poles is always present. Other characteristics, such as the relative extent of the cortex, may have some significance. Solereder and Meyer (1929) follow Nilsson in discriminating between ordinary roots and mechanical roots. This contrast is probably more one of degree than of kind. In the present study, the roots of Achlyphila disticha (fig. 7) might be sdid to be mechanical roots on account of their sclerenchymatous vascular core in which phloem is minimal. The presence of sclerenchyma in the core, with small scattered patches of phloem cells, suggests. such a designation for the roots of A. linearifolia (fig. 2) described below, but their difference from other roots is relatively minor. Likewise, the roots of A. sprucet and A. macrostachya might bear such a designation, and Solereder and Meyer place roots of the latter species in that category, along with A. longifolia. 1960 | ANATOMY OF GUAYANA XYRIDACEAE 67 Figs. 1-4. Portions of transections of roots of Abolboda, each with the vaseular core below and the cortex and exodermis above. Fig. 1. A. sprucei. K 166. Fia. 2. A. linearifolia. X 155. - Fig. 3. A. acaulis. K 245. Fie. 4. A. macrostachya var. macrostachya. * 143. 68 MEMOIRS’ OF THE NEW YORK BOTANICAL GARDEN [von. 10 The figure by Malme (1925) of A. vaginata shows much greater sclerification of the vascular core than any of the species in the present study, so that this species should probably be mentioned in this regard. In only a few instances (Abolboda acaulis, A. americana) were relatively large phloem groups, dis- tinctly alternate with xylem poles, observed. The roots studied here differ little in structure of the cortex, which consists of large cells associated with relatively small intercellular spaces. This condition is reported for Abolboda vaginata and the species of Xyris section Xyris by Solereder and Meyer. There are, in roots of Abolboda sprucei and A. americana, occasional groups of stellate parenchyma cells in the cortex. This condition is mentioned by Solereder and Meyer for Abolboda vaginata and a number of species of Xyris. The walls, especially the inner, of the hypodermis and sometimes the adjacent walls of the subhypodermal layer are thickened in all the species studied here (figs. 1,2,7). Very light thick- enings were noted in the hypodermal layer of A. americana and A. macrostachya var. macrostachya (fig. 4, yes right). Hypodermal thickenings, making pos- sible the designation of an ‘‘exodermis,’’ were noted by Solereder and Meyer in Xyris indica and X. lanata. Abolboda. 1. A. acaulis. Endodermal cells in this species (fig. 3) are very thick-walled, with a small lumen which is nearer the outer than the inner surface of the endodermis. The stele is tetrarch or pentarch; no central vessel is present. The pericyele is unbroken. 2. A. acicularis (fig. 6). The endodermis is composed of cells only moder- ately thicknened on all surfaces. The inner wall is very slightly thicker than the radial or inner wall. Monarch, tetrarch, and 8-arch conditions were observed ; in the last-named, a central vessel was present. Interruption of the pericycle by vessels is various, and both a single and all but a single vessel in contact with the endodermis were observed in different roots. 3. A. americana (fig. 5). In all the numerous roots examined, comprising several collections, the endodermis was composed of cells the inner walls of which were markedly thickened. This thickening tapers on the radial walls to rather thin at their juncture with the outer walls, on which no thickening is present. The following stelar conditions were observed: 2-, 3-, 4-, 5-, and 6-arch. All vessel& are in contact with the endodermis, with the occasional exception of one or two. 4. A. linearifolia (fig. 2). The walls of the endodermal cells are thickened in a U-shaped manner, but thickenings on the radial walls are tapered abruptly to their point of contact with the exterior walls. A ring of parenchyma cells with noticeable thickenings ensheaths the endodermis. The stele is 7- or 8-arch, with one or two central vessels. Phloem groups are small and scattered; cells of the stele other than vascular elements are thick-walled. The pericycle is not interrupted by vessels. ; 5. A. macrostachya var. macrostachya (fe. 4). The root observed possessed endodermal cells much more radially elongate than those of the preceding ones. The walls are markedly thickened, with the lumen quite eccentrically placed toward the exterior side of the cells. The stele is very large, and the root ob- served was 15-arch, with 15 central vessels. Phloem groups are small and scattered throughout the stele. The pericyele is not interrupted by vessels. 1960 | ANATOMY OF GUAYANA XYRIDACEAE 69 O°#Ce;' Pe ht ROS re e te oP" eg. pgccets Fig. 5. Abolboda americana, transection of vascular core of root. X 286. Fia. 6. Abolboda acicularis var. acicularis, transection of root, showing cortex and exodermis at right. «K 127. Fie. 7. Achlyphila disticha, transection of root showing cortex and exodermis at right. 157. Fig. 8. Achlyphila disticha, transection of rhizome, exterior above. < 70. 70 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 6. A. macrostachya var. robustior. The roots observed in this variety are similar to those of the above variety, but diver in the following respects. Two, rather than one, exodermis layers are present. The parenchyma sheath around the endodermis is two cells wide at points. The two roots observed were 13-arch with 18 central vessels and 15-arch with 16 central vessels respectively. The central vessels cannot be differentiated from the peripheral ones at all points. 7. A. sprucer (fig. 1). The cortex of roots of this species is quite distinctive in consisting of only two cell layers internal to the exodermis. The outer of these layers is composed of quite large cells, the inner of smaller cells which ensheath the endodermis. Plates of small arm-parenchyma cells interrupt this condition in places. The endodermis consists of cells thickened in a peculiar fashion, so that the lumen is turbinate as seen in transectional outline. Rarely, the endodermis is two-layered (two such instances are seen in figure 1) to the extent of one or two pairs of cells per stele. The root examined was 7-arch with a central vessel. The pericyele is not interrupted by peripheral vessels. Achlyphila. In respect of exodermis and cortex structure, the root of Achlyphila disticha (fig. 7) conforms to the pattern outlined above. The endodermis, which stains bright orange-red (rather than pink, as in Abolboda) with safranin, possesses U-shaped thickenings not exactly like any mentioned above. The endodermis may be two cells wide in places, although this condition is rare and localized. Two roots examined proved to be 11-arch and 13-arch respectively. No central vessels are present. This is surprising in view of the large diameter of the vascular core. Instead, one to several cells that have the wall-thickening and staining characteristics of endodermis cells (figure 7, dark area in center of vascular core) are present in this position. On account of their characteristics, they probably should be regarded as endodermal cells, despite their location. Phloem groups are extremely small and are scattered amone the sclerified cells of the vascular core. The pericycle is not interrupted. Significantly, there are two or three layers of cells (as compared to fewer in Abolboda) between the pericycle and the outermost vessels. Occlusion by tannin-like materials is char- acteristic of many vessels. Orectanthe. No roots of Orectanthe were available for study. Vessels. For each of the above species, serial longitudinal sections were prepared to show the presence of vessels. All the species described above proved to have true vessel elements with simple perforation plates. Vessels have been reported by Solereder and Meyer (1929) in roots of Xyris lacera, X. lanata, and X. indica. Likewise, Cheadle (1942) indicates vessels with simple perforation plates for roots of Xyris fleruosa and X. smalliana. Discussion. The species of Abolboda described here offer no features not previously mentioned by Solereder and Meyer for Abolboda or Xyris. The possibility that 1960] ANATOMY OF GUAYANA XYRIDACEAE 71 endodermis-cell characteristics could be used as specific characteristics, sug- gested by Malme in 1925, appears to be valid. Malme shows very thick-walled endodermal cells with centrally placed lumina for A. pulchella and A. vaginata. He reports prominent thickenings on the inner wall, tapering sharply on the radial walls, and absent on the outer walls, for endodermal ¢ells in roots of A. poeppigu, A. grandis, and A. macrostachya. Malme’s figure for A. macro- stachya is markedly different from the condition illustrated here for that species. The endodermal cell thickenings Malme reports for A. poarchon and A. abbre- viata are like those illustrated in the present study for A. acicularis. Interestingly, Malme’s figures all show an uninterrupted pericycle (A. vagi- nata, A. macrostachya, A. poarchon). The species of Abolboda in the present study in which uninterrupted pericycle was observed all belong to the group termed ‘‘larger-stemmed abolbodas’’ in sections below. Malme’s figures also suggest a larger number (six or more) of xylem poles, a feature also character- istic of the group just named. Thus as Malme (1925) claims, different types of root anatomy in Abolboda appear to be characteristic of species or species- eroups; from the example of A. sprucei described above, one may surmise that such characters are not always limited to the stele, but may be found in the cortical zone as well. The several layers of cells between endodermis and outermost vessels. the absence of central vessels, the presence of endodermis-like cells in their place, and the distinctively staining endodermis thickenings are characteristics which, taken together, would seem to give the roots of Achlyphila a generic differentia- tion from the other three genera. In fact, some of these characteristics seem new to Xyridaceae, although the sum of characteristics in this genus would not exclude it from the family. On the basis of the present study, one may conclude that the roots of Abolboda offer no anatomical features (other than greater size, and therefore more numerous vessels) which cannot also be found in Xyris roots, as the account of Solereder and Meyer (1929) illustrates. STEM As the summary of Solereder and Meyer shows, remarkably little is known about stem anatomy in Xyridaceae. The only facts that have been established are that bundles are amphivasal, individual bundles may be sheathed with sclerenchyma (incompletely in X. Janata), and that (in X. lanata) the bundles may form concentric bands. As an additional generalization, the writer would like to add the fact that in all taxa he examined, vascular bundles tend to be more or less amphivasal except where they are demarcated as leaf traces, in which case the structure of the bundles tends to be more nearly collateral. The data given below are derived from study of mature portions of rosette stems, except for Achlyphila, which has a rhizomatous habit. At and near the base of a stem, especially in the smaller-stemmed species of Abolboda, departing roots may be numerous in the cortex, and the vascular bundles may form a very small group in the center of the stem. In A. americana, no part of the stem (except the very youngest) was free from departing roots. The types of stem anatomy observed in the XY yridaceae under study here fall into four main groups. LY 72 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Smaller-stemmed Abolbodas. The four species, A. acaulis (fig. 9), A. acicularis (fig. 10), A. americana, and A. ciliata (fig. 11), that are considered under this category show four features: (a) the bundles other than those that are leaf traces are most often enclosed in a sclerenchyma cylinder; (b) sclerenchyma ensheaths departing leaf traces indi- vidually; (c) the ground tissue of the stem (other than sclerenchyma) consists of more or less spongy tissue lacking in starch; and (d) the bundles (except leaf traces) are present in a cylinder and no bundles are present in the central portion of the stem. The variations on this pattern are as follows. 1. A. acaulis (fig. 9). In this species, departing leaf traces are most often incompletely sheathed with sclerenchyma; such sclereids are few or absent at the outer (phloem) pole. Centrally placed in the core of parenchyma is a nest of sclereids. This nest of sclereids occurs along the length of the stem, and is present even in the most basal region, where other sclerenchyma is wholly lacking. 2. A. acicularis (fig. 10). The central portion of the stem, including bundles other than leaf traces, is sclerified. Sclerenchyma forms a conspicuous sheath around leaf traces; it is more abundant at the xylem pole of each bundle. The bundles other than leaf traces form a single ring. Sclerenchyma may form a cylinder, as illustrated, may occupy the entire central portion of the stem, or may, as illustrated for A. acaulis, form a nest within the ring of bundles. The latter condition occurs in the base of the stem, and the cylinder is characteristic of the upper portion of a stem. 3. A. americana. Sclerenchyma (other than that which forms a cylinder in the center of the stem) is present only at the xylem pole of departing leaf traces. 4. A. ciliata (fig. 11). This species shows essentially the same features as A. americana, although the stem is somewhat larger. . Larger-stemmed Abolbodas. This group includes A. linearifolia (fig. 12), A. macrostachya and its vari- eties (fig. 13, var. angustior), and A. spruce (fig. 14). These species agree in possessing (a) much wider stems than species of the above group; (b) no scleren- echyma other than individual sheaths on bundles; (c) compact, never spongy ground tissue, often differentiated into a starch-poor zone outside and an inner core (figs. 12, 13, inside dotted lines) which contains numerous elongate starch orains; "and (d) bundles throughout the central portion of the stem, more mvrerous in a eylinder just inside the outer limit of starch-rich tissue. In all preparations, the ground tissue consists of more or less isodiametric cells asso- ciated with relatively small intercellular spaces. Starch may be expected to vary in amount depending on the state of development of the plant. Variations with respect to species are as follows. 1. A. linearifolia (fig. 12). Many of the bundles lack sclerenchyma. Where present, it takes the form of a semicircle at the interior pole of the bundle. Sclerenchyma is associated with the larger bundles, and may be an indication of major bundles which enter leaves a short distance above the level of sectioning. 2. A. macrostachya. In A. macrostachya var, macrostachya and var. robus- tior, the same condition described for A. linearifolia was observed. In A. macros- tachya var. angustior (fig. 13), however, all bundles were found to be associated with sclereids, which may be quite abundant near some bundles. Some leaf traces are completely ensheathed by such sclerenchyma. 1960 | ANATOMY OF GUAYANA XYRIDACEAE Figs. 9-14. Portions of transections of rosette stems of Abolboda, exterior above, pat terned to show distribution of various tissues. Fic. 9. A. acaulis. ciliata. Fic. 12. A. linearifolia. Fic. 13. A. macrostachya var. angustior. Fic. 14. AL sprucei. Broken line in figures 12, 13 delimits the outer starch-poor ‘region from the inner starch-rich zone. Patterns as follows: stippled = sclerenchyma; white inside bundles phloem; black = divisions. xylem; parenchymatous ground tissue left bank. Scale = 2 mm. in 1 mm. Fig. 10. A. acicularis var. acicularis; leaf base in paradermal section, above; endogenous root at right: Fie. 11. A. 74 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 1960] ANATOMY OF GUAYANA XYRIDACEAE i) 3. A. sprucei (fig. 14). All bundles are associated with abundant scleren- chyma sheaths, with few exceptions. Such sclerenchyma may in fact incorporate several bundles. Bundles that lack sheaths are frequently transversely oriented and tend to delimit a cortical zone, in which leaf traces are present, from the central zone. Orectanthe. As might be expected, the large stems of Orectanthe have an anatomy quite distinct from that of the above species. There is a difference between the pattern found in O. ptaritepuiana (fig. 15) and that which was observed in O. sceptrum (fig. 16). In O. ptaritepwiana, a cortical zone is clearly delimited by a scleren- chymatous cylinder which consists of lignified parenchyma cells. The inner margins of this cylinder are irregular, for some bundles and their sclerenchyma- tous sheaths are joined to it and others are independent in the thin-walled central ground tissue. Outside of the sclerenchyma cylinder, the parenchyma is also thin-walled. Bundles in this region are smaller than those within the cylinder, and are either lacking in sclerenchyma sheaths or very sparingly pro- vided with such tissue. In O. sceptrum (fig. 16) the entire central portion of the stem consists of lignified ground tissue. Wall thickenings are much greater toward the exterior of the sclerenchymatous core. In addition, wall thickenings are greater on the 2—3 layers of cells that sheath bundles in the sclerenchymatous zone. The cortical region consists of non-lignified parenchyma which contains vascular bundles smaller than those encased in the seclerenchyma core. These leaf traces are ordinarily provided with fibrous sheaths which completely encase them. Achlyphila. In the single species, prostrate rhizomatous stems are present. Upright stems are formed in connection with production of inflorescences, and these are con- sidered under ‘‘Inflorescence Axis’’ below. The stem described here, therefore, is the rhizome, which differs from rosette stems of Xyris, Abolboda, and Orec- tanthe in its long internodes. This rhizome (fig. 8) possesses a marked eylinder of thick-walled sclereids which delimits the cortical region of thin-walled paren- chyma cells. The outermost layer of this cylinder consists of cells that are, in their thickening patterns and staining properties, very similar to root endo- dermis cells. The occurrence of endodermis in rhizomes of monocotyledons is not uncommon, and these cells should be so interpreted here. The inner margins of the sclerenchyma cylinder are irregular in outline. In the central portion of Figs. 15, 16. Portions of transections of rosette stems, of Orectanthe, exterior at right. Fie. 15. O. ptaritepwiana. Fic. 16. O. sceptrum. Seale for figures 15 and 16 = 2 mm in 1 mm divisions. Figs. 17-20. Portions of transections of inflorescence-axis bracts, exterior at right. Fic. 17. Abolboda ciliata, section midway along length of bract. Fig. 18. A. macrostachya var. macrostachya, section from tip of bract. Fic. 19. Same, section midway along length of bract. Fic. 20. A. macrostachya var. robustior, section midway along length of bract. Fie. 21. Orectanthe sceptrum, transection of a flattened sepal. Fies. 22-24. Transections of bracts of inflorescence proper, taken midway along length of bract, exterior face above. Fic. 22. Abolboda macrostachya var. robustior. Fic. 23. A. linearifolia. Fa. 24. A. americana. Seale for figures 17-24 = 2 mm in 1 mm divisions. Patterns as follows: fine stippling = thick-walled sclerenchyma. coarser stippling = thin-walled sclerenchyma. cross-hatched = chlorenchyma; white in bundles = phloem; black = xylem; parenchymatous ground tissue left blank. ys A]. SS a Lee i oh 2 My Cao 20 e ("4 Yo ¢ AL? A sets & 9 ity ae } BERR oo ee SPO) St acer) O gers, ty MG OO as Oe: Bee Ged & eeessarees, ee Edie MRR autho 9 > a t. ra J LAG anti 0° sey ry RE Se IU Or ES 3 SARE, ah, osty2 Scene AG Sip te MeGomevance Ts Wit HDMI ROL ne TNANIDOUIOSSOOS Figs. 25-30. Transections of leaves, taken approximately midway along length of leaf, adaxial face above. Fic. 26. Abolboda americana, Fia. 26. A. acicularis var. acicularis. @) “hs © 1960} ANATOMY OF GUAYANA XYRIDACEAE ue the stem, bundles are embedded in thin-walled parenchyma. Each bundle 1n this region is completely ensheathed by lignified cells like those of the sclerenchyma eylinder. Leaf traces are uncommon in the cortical region (on account of the long internodes), but where these are present, each bears a sclerenchyma cap on the xylem face of the bundle. There are occasionally a few thick-walled lignified parenchyma cells in the cortical region. Discussion. Despite the fact that the meager literature on stem anatomy does not suggest the usefulness of this organ for systematic purposes, excellent generic and specific characteristics are seemingly present, as outlined above. The smaller- stemmed species of Abolboda are distinctive in their lack of central bundles, their sclerenchyma cylinder, and their spongier, starch-free parenchyma, as compared to the larger-stemmed species. The two species of Orectanthe are like the larger-stemmed species of Abolboda in bundle distribution, but show a promi- nent lignification of parenchyma; this lignified core is clearly delimited from a thin-walled cortical region. The rhizomatous stem of Achlyphila is distinctive in Xyridaceae, but is not extremely unlike stems of Abolboda in its construc- tion. Features of generic value include its prominent sclerification, distribution of bundles, and development of an endodermis. Vessels. Longitudinal sections of stems in all the taxa named above were prepared, and these were used to demonstrate that in all taxa named, vessel elements with simple perforation plates occur. Cheadle (1942) has indicated such vessel ele- ments for two species of Xyris. LEAF The same groupings used for descriptions of stem anatomy may be used here. The species and species-groups prove highly diversified, so that few generaliza- tions can be offered. Smaller-stemmed Abolbodas. Species in this group are distinctive in their smaller leaf size, the relative lack, or restriction of, photosynthetic tissue, and the presence of longitudinally- oriented pockets of parenchyma which collapses at maturity and appears to have been composed, in some instances at least. of chlorenchyma cells. Hypodermis is absent, or present only in the more apical portions of the leaf. Stomata are restricted to the lower surface. 1. A. acaulis (figs. 31, 35). The basal portion of a leaf (fig. 31) consists of parenchyma, isodiametrie as seen in transectional outline, in which 4-5 veins are embedded. Between the veins, zones of collapsed parenchyma are present. Selerenchyma completely ensheaths the veins; such sheaths are usually two cells in thickness. At a higher level (fig. 36) a hypodermis begins to be differen- tiated; it is prominent (by virtue of contrast with chlorenchyma) only in the most apical portions of the leaf, where it consists of gelatinous-walled sclereids, Fig. 27. A. ciliata. Fic. 28. A. macrostachya var. robustior. Fic. 29. Orectanthe ptarite- puiana. Fig. 30. O. sceptrum subsp. occidentalis. Scale = 0.5 mm in 0.1 mm divisions. ‘ 78 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 at least in places. In the section shown (fig. 36) three main veins, with fibrous sheaths, are present, and chloroplasts are present in the mesophyll parenchyma, although they are rather sparse. 2. A. acicularis (fig. 26). Leaves of this species are similar to those of A. acaulis. A section approximately halfway along the length of the leaf (fig. 26) reveals three veins, each surrounded by sclerenchyma and embedded in chloroplast-free parenchyma cells isodiametric in transaction. Cells rich in chloroplasts form a layer immediately beneath the epidermis on both surfaces. Between the veins, pockets of collapsed parenchyma cells occur. A thick-walled hypodermis occurs in the most apical portion of the leaf, where the mesophyll is composed of spherical chlorenchyma. 3. A. americana (fig. 25). This leaf shows little differentiation from base to apex, except for reduction in number of veins. Especially prominent are the longitudinal zones of collapsed parenchyma. Some of the smaller bundles lie in these zones. All of the bundles are completely ensheathed by sclerenchyma. The parenchyma is composed of cells, isodiametric in transection, with relatively few chloroplasts. These cells may be weakly differentiated as a hypodermis in the adaxial portion of the leaf. The leaves of A. americana are certainly within the range of variation of Abolboda, but an additional similarity may be noted to leaves figured by Arber (1925) for Eriocaulon septangulare. 4. A. ciliata (figs. 27, 35). An extensive series of transections of a leaf of this species showed the nature of differences, characteristic of A. ciliata, between base and apex. The most basal portion (leaf sheath) consists only of thin-walled parenchyma in which approximately seven bundles, without fibrous sheaths, are embedded. In the upper portion of the leaf sheath, sclerenchymatous sheaths are present and collapsed parenchyma cells occur between veins, resembling the condition shown for A. acaulis in figure 31. The margins of the leaf at this level are biseriate. Segments of this wing split off, forming the ‘‘cilia’’ implied in the species name. Above the level of the sheath, (fig. 27), the leaf is markedly thicker, lacks tapered margins, and has a hypodermis, one or two eells thick, on the adaxial face of the leaf. The mesophyll (other than the pockets of col- lapsed cells) consists of spherical cells with numerous chloroplasts. Seven bundles, each surrounded by a sclerified sheath, are present at this level. At higher levels, such as that shown in figure 35, the outermost bundles are not present. «At the margins of the leaves, a portion of the hypodermis, which ex- tends around the edges of the leaf toward the abaxial surface, consists of sclerified cells. At this uppermost level, the chlorenchyma cells are differen- tiated (mostly by relative size of intercellular spaces) into palisade and spongy tissue. The level shown in fig. 35 1s notable for the greater thickness of the leaf, a thickness due not only to the greater number of hypodermis layers, but to the greater number of chlorenchyma layers as well. Larger-stemmed Abolbodas. . Despite certain distinctive features in the leaf of A. linearifolia, there is agreement among A. linearifolia, A. macrostachya, and A. sprucei in the follow- ing respects: leaves thicker and larger than those of the above group; hypo- dermis present, consisting of 3-5 layers of cells; parenchyma like that of the hypodermis (polygonal in transection, markedly elongate in longitudinal sec- tion) also present and bundle sheaths (surrounding fibrous bundle-sheath layers) 1960| ANATOMY OF GUAYANA XYRIDACEAE 79 Figs. 31-33. Transections of leaves of Abolboda, adaxial face above. Fic. 31. A. acaulis, section taken toward base of leaf. * 80. Fic. 32. A. linearifolia, section taken toward base of leaf. & 65. Fic. 33. The same, section taken midway along length of leaf; margin at right. < 65, | ‘ . 80 MEMOIRS, OF THE NEW YORK BOTANICAL GARDEN [vou. 10 and bundle-sheath extensions; the remainder of the parenchyma being either of globular chlorenchyma cells or collapsed parenchyma cells. 1. A. linearsfolia (figs. 32, 33). In the basal portion of the leaf (fig. 32) there is a large number of vascular bundles. These alternate with pockets of collapsed tissue. Most significant is the fact that veins occur as bundle-groups. Most of the veins are thus compound, and consist of 2-3 bundles. This condi- tion is unique in Abolboda, and proves an exception to the statement of Solereder and Meyer (1929) that the bundle-group type of construction in Xyridaceae is restricted to XYyris. Each bundle in the bundle-groups is surrounded by a fibrous sheath which is fused with the sheaths of neighboring bundles in the group. At a higher level (fig. 33) the same type of bundle is evident. There are, as described for A. ciliata, marginal hypodermal fibers, and the hypodermis other than this consists of about five cell layers. Bundle-sheath extensions are present, but relatively infrequent—only 1-3 may be present in a section at this level. The remaining mesophyll consists of spherical chlorenchyma cells which occur beside the hypodermis, lower epidermis, and around the bundle sheaths. Meso- phyll not adjacent to these areas consists of collapsed parenchyma cells. The structure of leaf-tips is considered below in a separate section. 2. A. macrostachya. The sheathing portion of the leaf of A. macrostachya var. macrostachya (fig. 34) is like that described for A. linearifolia in the nature of chlorophyll-free and collapsed parenchyma distribution. It is thinner, how- ever, and the bundles, which are not grouped, possess massive sclerenchyma bundle sheaths. At upper levels in the leaf of A. macrostachya var. robustior (fig. 28) a hypodermis of 3-4 layers occurs. Bundle sheath extensions consist of cells similar to the hypodermis and are present on all but the smallest veins. The remaining mesophyll consists of chlorenchyma, more densely arranged to- ward the adaxial surface. Only the larger veins have lignified elements in the bundle sheath; these are more abundant on the abaxial faces of the bundles. In comparable portions of the lamina of A. macrostachya var. macrostachya and var. angustior, which are much alike in leaf anatomy, only 1-3 bundle sheath extensions are present. Fibrous bundle sheaths generally are complete in var. angustior, whereas they often are present only along the abaxial surface of veins in var. macrostachya. In other respects the three varieties are alike. Structure of the leaf tips is described below. 3. A. sprucei (fig. 37). The sheath portion of the leaf in this species is like that shown for A. macrostachya (fig. 34) and requires no further comment. The upper portion of the lamina in A. sprucei (fig. 37) also shows similarity to that of A. macrostachya. A hypodermis 2-5 cell layers wide is present, and no marginal sclerenchyma occurs. The remaining mesophyll consists of globular chlorenchyma cells which are more densely arranged beneath the hypodermis. The bundle sheath of all veins, in addition to the outer layer of chlorophyll-free parenchyma, consists of a massive fibrous sheath. 4. Leaf-tip. The three species above are alike in having a thorn-like ter- minus which occupies the 1-2 mm at the leaf apex. This type of apex in itself is not exceptional, nor is the fact that xylem alone without phloem composes the terminal portion of the vein, for this condition has been reported in other monocots (Arber, 1925). The tracheids in these species (fig. 41) tend to be separated from each other by parenchyma cells. The genuinely remarkable feature concerning these tracheids is the presence of very wide bands which 1960 | ANATOMY OF GUAYANA XYRIDACEAE 81 mye af Sse tx + @ *. Fy rd *e 2 & Fie. 34. Abolboda macrostachya var. macrostachya, transection of leaf base, adaxial face at right. X 74. Fic. 35. A. ciliata, transection of upper portion of leaf, adaxial face at right; margin below. X 90. Fic. 36. A. aceulis, transection of upper portion of leaf; margin at left, adaxial face above. K 123. Fig. 37. A. sprucei, portion of transection taken midway along length of leaf; margin at right, adaxial face above. X 108. Fic. 38. A. sprucei, transection of ovary crest. * 108. , 82 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 form the helical wall thickenings (fig. 41, enlarged in fig. 40). These bands, which are homologous with the typical helical or annular bands on protoxylem tracheids, are up to half the lumen in width. In helical tracheids more than one helix (usually two) may be present per tracheid. The bands are thicker toward the center of the cell, reflecting a ‘‘bordered’’ condition, as seen in figure 40, left. As seen in a transection of the leaf tip (fig. 42), the tracheids appear like fibers, because the wide bands resemble the thick walls of fibers. There seems little doubt that these represent specializations of ordinary annular and helical protoxylem tracheids. The transition to them from normal tracheids is abrupt, but elements with transitional width of thickening bands may in fact be found. The elements with the widest bands occur at the most apical portion of the leaf tip. The function of these peculiar tracheids is difficult to imagine. There is no hydathode formation in the terminal portion of the leaf (which is covered with one to several layers of hypodermal sclereids), nor any stomata in the terminus, so that this function may be ruled out. Although these tracheids are probably rare in angiosperms at large, they have been described in Cactaceae, as the account of that family in Metcalfe and Chalk’s Anatomy of the dicoty- ledons shows (Cf. also A. Brongniart, Arch. Mus. Hist. Nat. Paris 1: 405-461. 1839). Orectanthe. Leaves of Orectanthe are basically not unlike those of the species of Abolboda just described, but differ from them in having a broad, flat shape, and an abaxial hypodermis (in some places, at least) as well as an adaxial one. The chlorenchyma cells are elongate in longitudinal section, and have symmetrically placed short arms (fig. 43). They correspond to the cells shown for Xyris lanata by Solereder and Meyer (1929, fig. 11b). 1. O. ptaritepuiana (fig. 29). In this species, 3-4 nell layers of hypodermis are present on the adaxial surface. The same type of parenchyma cells which comprise the hypodermis are present as a sheath extension on the larger veins, and as a single layer of hypodermis on the abaxial surface between the bundle- sheath extensions (at least in some places). The remaining portions of the mesophyll are pockets, oriented longitudinally in the leaf, which contain a uni- form tissue composed of the peculiar chlorenchyma cells described above. The smaller bundles which do not have sheath extensions are embedded in this tissue. The bundles are elongate in transectional outline, and are completely ensheathed by a sclerified bundle sheath. These bundles are strictly collateral in structure. 2. O. sceptrum. Both typical O. sceptrum and subsp. occidentalis (fig. 30) are alike in having leaves thinner than those of O. ptaritepuwiana. They are like leaves of O. ptaritepuiana in all details of anatomy except that (1) they often have an abaxial hypodermis nearly as wide as the adaxial one, (2) they have a prominently selerified epidermis, and (3) the bundles are round, not elongate, in outline. The thickenings on the walls of the epidermis are narrowest on the outer wall. In upper levels of the leaf of O. sceptrum subsp. occidentalis, the parenchyma of the hypodermis and bundle-sheath extension was found to be more thick-walled and lignified. A section of the sheath portion of the leaf in this subspecies reveals that the structure of this portion of the leaf is identical with that of the leaves of the larger-stemmed abolbodas. CO OW 1960 | ANATOMY OF GUAYANA XYRIDACEAE Fig. 39. Achlyphila disticha, portion of sagittal section of upper leaf, showing sclereids which comprise the epidermis. * 200. Fig. 40. Abolboda sprucei, enlarged portion of a section also shown in figure 41; at left, a tracheid with very wide annular thickenings; in center, a parenchyma e¢ell; at right, out of focus, wide band from a_helically-thickened tracheid. K 915. Fic. 41. The same, showing a larger area of a sagittal section of the thorn-like tip of a leaf. Portions of tracheids (with very thin intervening walls) are shown, right, and left, and in the center (dark) parenchyma cells. The tracheids, left, have annular bands; those at right shown annular bands above and helical bands below. &* 523. Fia. 42. A. macrostachya var. macrostachya, transection of thorn-like leaf tip, showing wide-helix and wide-annular band tracheids interspersed among parenchyma cells. & 285. Fic. 43. Orectanthe ptaritepuiana, portion of parasagittal leaf section, showing abaxial epidermis (above) and about three layers of arm-palisade cells. & 238. [vou. 10 YORK BOTANICAL GARDEN NEW ri] 4 TEMOIRS OF THE \ 84 rie ea i re a t ty Af $9, Ad | 44. Half of transection of upright stem (inflores- Fig. 45. Portion of transection of sheating base, adaxial surface at 46. Transection of leaf base, at higher level than figure 45, showing Fig. Achlyphila disticha. 31. —4.7. Figs. 44 x 1xis ). © c cence * 100. Jie: right. 1960] ANATOMY OF GUAYANA XYRIDACEAE 85 Achlyphila. * Corresponding with the equitant habit of leaves, Achlyphila disticha shows anatomical features unusual or unique in Xyridaceae. Among these features are the sclerified epidermis, the bands of sclerenchyma in the leaf sheath, and the central zone of thick-walled parenchyma in the upper portions of the leaf. A transection of the sheathing portion of the leaf (fig. 45) shows three distinct zones of mesophyll: the outermost consists of spherical chlorenchyma cells; interior to this is a band of sclerenchyma in which the bundles he. This scleren- chyma zone represents the widening, and fusion, of the individual sclerenchyma- tous bundle sheaths, which are separate at higher levels (figs. 46, 47) in the leaf. The innermost layers, present only on the adaxial surface of the central leaf sheath (fig. 46), consist of thick-walled parenchyma cells, isodiametric and polygonal in transection. Because of the equitant habit, sections at higher levels show the fusion of the two halves (fig. 46) into a conformation elliptical in transection (fig. 46). Here the bundles and their sheaths are separate, with no intervening sclerenchyma. The chlorenchyma ean be differentiated into palisade and spongy at this level. The cells at the dorsal and ventral edges of the leaf (sclerified epidermis) are prominently elongate as seen in transection, forming irregular shapes. As seen in longitudinal section (fig. 39), the adjacent edges of epidermal calls are often raised into emergences, whereas the central surface of such cells is depressed. This configuration of epidermal cells is re- ported by Solereder and Meyer (1929) for several Xyris species and figured for X. montivaga. The elongate sclerified epidermal cells, as seen in transection, closely match those which Arber (1925) figures for X. anceps, although this species, unlike XY. montivaga, does not have the undulate contours of epidermal cells as seen in longitudinal section. Sections of the leaf of Achlyphila at the level shown in figure 47 show that the central parenchyma of the leaf is thick- walled, probably lignified, and has an elliptical shape. At higher levels, there is a diminution both in the area of the central parenchyma and the number of bundles which surround it. Discussion. Malme’s claim in 1925 that leaf anatomy is rather stereotyped in Abolboda as compared to Xyris, except for the number of veins, does not appear to be justified on the basis of the present study. A number of seemingly excellent specific and species-group characteristics have been described above. Leaf anatomy of Orectanthe reveals the similarity of that genus to Abolboda as well as the generic distinctions of the two species. The leaf of Achlyphila is not unlike those of other Xyridaceae in its general structure. For example, Xyris asperata has a similar equitant habit (although the anatomy is quite different), as illustrated by Arber (1925). The presence of a ring of bundles around a zone of thick-walled parenchyma seems unique in the family, however. The epidermal characteristics although quite unlike those of Abolboda and Orectanthe, find parallels in Xyris. portions of the two wings. * 70. Fic. 47. Transection of upper (equitant) portion of leaf, showing slightly more than half; orientation as in figure 46—morphologiecally adaxial por- tion of leaf above. X 80. . 86 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Vessels. Although Solereder and Meyer (1929) do not mention the occurrence of vessel elements in leaves of Xyridaceae, longitudinal sections of leaves revealed vessel elements with simple perforation plates in the following species: Abol- boda linearifolia, A. macrostachya var. angustior, A. macrostachya var. macro- stachya, and A. macrostachya var. robustior. Vessels, therefore, definitely do occur in leaves of at least the larger-leaved species of Abolboda. Vessels with simple perforation plates are indicated by Cheadle (1942) for leaves of Xyris flexuosa and X. smalliana. INFLORESCENCE AXIS Solereder and Meyer (1929) have concluded that Xyris differs from A bolboda in the general structure of the inflorescence axis. Xyris, according to these authors, possesses bundles embedded in (rarely outside of) the sclerenchyma ring, whereas Abolboda has bundles scattered through the central ground tissue (‘‘pith’’) as well. This contrast appears to be justified on the basis of the present study. Solereder and Meyer also claim that bundles of the graminean type are absent in the inflorescence axes of Abolboda although they are present in Xyris. I have found that the more basal portions of inflorescence stems in Abolboda tend to have ‘‘ordinary’’ collateral bundles, or ones which are some- what amphivasal. At higher levels, the graminean-type bundles were seen in ‘“nith’’ of all the taxa considered here except Abolboda acicularis, A. americana, and Achlyphila disticha. Abolboda. Variables within a single inflorescence axis of Abolboda include the relative presence of photosynthetic tissue and the thickness of the sclerenchyma ring. Chlorenchyma appears to be absent from those portions of the axis most distal and most proximal to the base of the plant. Chlorenchyma, where present, con- sists of nearly spherical cells; some may be somewhat elongate radially, and form a palisade. If chlorenchyma is not present, parenchyma cells polygonal in outline in transection (elongate in longitudinal section) extend from the epidermis to the sclerenchyma ring. If chlorenchyma is present, a single layer of non-photosynthetic parenchyma is often present outside the sclerenchyma ring. If chlorenchyma is present, a single layer of non-photosynthetic par- enchyma is often present outside the sclerenchyma ring, particularly at those points where bundles are adjacent to the outer face of that ring. The scleren- chyma ring consists of several layers of thick-walled lignified cells. The outer- most may have much thicker walls than layers internal to it. 1. A. acaulis. Because of the nearly sessile habit of the inflorescence in this species, no structure comparable to the inflorescence axis in the other species can be studied. 2. A. acicularis. This species (fig. 49) shows a number of features typical for the genus. Within the epidermis, both palisade and spongy palisade chlorenchyma occur. The single layer of non-photosynthetic parenchyma out- side the sclerenchyma ring is ordinarily present. The outer face of each of the bundles (about 7) which occur outside the ring is sheathed by a single layer of sclerenchyma. No bundles occur within the sclerenchyma ring itself, 1960 | ANATOMY OF GUAYANA XYRIDACEAE 87 which is about three layers thick. A number of scattered bundles are present in the ‘‘pith’’ region. 3. A. americana (fig. 48). The cortical region of the inflorescence stem is notable for the occurrence of pockets (which run longitudinally in the exis) of collapsed parenchyma cells—like those which occur in a similar manner in the leaves of this species. The collapsed parenchyma pockets occur alternately with bundles of the outer circle. The remainder of the cortical region consists of globular parenchyma cells. The bundles adjacent to the outer margin of TK ONSUG dA HER PERT 0s pore ts bam a Ce Be BEG Faye te ios OS ogra SS FARE ike ame cee ae rg iiuemict aston CON Nd mAer**— a S' Cf) BEY RARE SCY ety) COs s ce@ Oh: Orda y Oe Sean {) re eK ae. c , as oe < rf a: iu Figs.’ 48-52. Portions of transeetions of inflorescence axes of Abolboda. exterior above. Fig. 48. A. americana. Fig. 49. A. acicularis var. acicularis. Fie. 50. J ee A 0 AREY ae Oeene Oc ys LR N 4 ANY SoH a al Soszu : LSA ARE ST SSO SISOS gs DACINTOO ESS OB RE TWN SCNOYS e ies GQ we i Cee Qe @ 7 ©, /> 6) aes es wie Lk ald Cs ee a see 7 eee Ser SO): @)2s) [OYRS @® oe. es Oo: SOeS gs sine aa : Vy; anna Sem 0) Re © Figs. 53-55. Transections of inflorescence axes, exterior at right. Fig. 53. Abolboda macrostachya var. macrostachya. Fig. 54. A. macrostachya var. robustior. Fie. 55. Orect- anthe sceptrum. Scale = 0.5 mm in 0.1 mm subdivisions. ; \ + 90 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN ivou. 10 9. A. sprucei (fig. 52). The inflorescence stem of this species shows no features not indicated above, but illustrates the generic characters rather clearly. The cortical zone of this axis is composed of spherical chlorenchyma cells. Exterior to the sclerenchyma ring is a circle of bundles; this circle is enveloped by a layer of non-photosynthetic parenchyma. The outermost layer of the scelerenchyma ring is very thick walled, and several less prominently sclerified cell layers occur internal to it. All the central bundles occur in the thin-walled ground tissue, although some are adjacent to the sclerenchyma ring. Bundles are of the graminean type or transitional to that type from the ordinary collateral. Orectanthe. An inflorescence axis of only one species, O. sceptrum (fig. 55), was see- tioned. This stem showed a number of noteworthy features. A sclerified epidermis is present. Beneath this are about two cell-layers differentiated as a hypodermis. The remainder of the cortical zone consists of spongy chlorenchyma. Within the chlorenchyma are a number of bundles, most of which have a sclerenchy- matous sheath. The sclerenchyma ring contains a number of bundles, not dis- posed in concentric circles. The thin-walled ground tissue intergrades into the sclerenchyma ring. Bundles of the sclerenchyma ring and central ground tissue are intermediate between amphivasal and collateral patterns. Achlyphila. The markedly flattened inflorescence axis of this genus (fig. 44) suggested that more or less prominent differences from Abolboda would,be present. In fact, despite the obvious differences in shape, the similarities are perceptible. To be sure, the epidermis (which is two-layered in places at the edges of the axis) is heavily sclerified. Then, too, bundles (each with a fibrous sheath) occur in the cortical region on each edge of the axis. Aside from these differences, one can cite similarities in the chlorenchyma: two layers of palisade and several spongy layers are present. Adjacent to the outer margin of the scle- renchyma ring—or ‘‘ellipse’’—bundles are present, each sheathed by a single layer of fibers along the outer surface, as in Abolboda. Bundles are present in the sclerenéhyma, toward the inner margins of its extent. The central ground tissue consists of thin-walled parenchyma and (unlike Abolboda, but like Xyris) contains no bundles. The last-cited feature suggests that the resemblances of Achlyphila may lie closer to Xyris than to Abolboda. Because Achlyphila is unique among Xyridaceae in not having a congested inflorescence, elongate pedicels are present. As is shown in figure 82, the pedicel is triangular in outline. Like the inflorescence axis, the pedicel has a sclerified - epidermis. Two circles of bundles. one adjacent to the inner margin of the sclerenchyma ring (or, at lower levels, just within the ring) may be found. A™ central zone of thin-walled parenchyma is present. Discussion. The above descriptions must not be construed as exhibiting the full range of variation either in a species or in an individual axis, although an attempt is made to suggest the nature of such variation. Because of this variability, 1960] ANATOMY OF GUAYANA XYRIDACEAE 91 species characters are difficult to offer. The inflorescence stems of Abolboda macrostachya are distinctive on account of their greater size (with more numer- ous bundles). Special features in the axes of A. americana (a single circle of ‘‘pith’’ bundles; collapsed parenchyma pockets) and A. ciliata (1-2 circles of ‘‘pith’’ bundles; prominent protoxylem lacunae in these) may be noted. The extent of sclerenchyma seems roughly correlated, in Abolboda, with size of the axis, and the sclerenchyma ring in A. macrostachya is much wider than in A. americana. Likewise, the very wide sclerenchyma cylinder in Orectanthe may be correlated with the large diameter of that axis. The sclerified epidermis, the lack of concentric arrangement of bundles, the presence of scattered bundles in the cortical region, and the presence on these of fibrous sheaths are all characters which differ to a certain extent from comparable conditions in Abolboda, and enforce the generic status of Orectanthe. The descriptions of Achlyphila above have shown that inflorescence-axis anatomy serves both to distinguish this genus generically, and to relate the genus to other Vyridaceae, probably to Xyris most closely. Vessels. Solereder and Meyer (1929) cite a report of vessels with simple perforation plates in inflorescence axes of NXyris caroliniana, X. indica, and X. lacera. In the present study, such vessel elements were observed in longitudinal sec- tions of inflorescence axes of the following taxa: Abolboda acicularis, A. ameri- cana, A. macrostachya var. macrostachya, A. macrostachya var. robustior, and Orectanthe sceptrum. Cheadle (1942) indicates vessels with simple perforation plates in XYyris fleruosa and X. smalliana. INFLORESCENCE BRACTS One or more bracts tend to occur along the inflorescence axis, and several are associated with the congested inflorescence itself. The former are here termed inflorescence-axis bracts and the latter inflorescence bracts. These bracts Show patterns of structure different from those of sepals or leaves, and are worthy of description. In some instances (Abolboda acicularis, A. ciliata) the difference between structure of inflorescence-axis bracts and sepals is not very great. Because inflorescence-axis bracts and inflorescence bracts vary in structure from base to apex, and even within a plant, depending on their position, their structure is difficult to categorize accurately in all instances. Sections studied are discussed by species. Abolboda. 1. A. acaulis. Because of the subsessile nature of the inflorescence in this species, bracts might best be regarded as inflorescence bracts. Such bracts have three veins in a non-photosynthetic parenchyma. Sclerenchyma is absent. 2. A. americana (fig. 24). An inflorescence bract is illustrated for this species. Three veins are present. The middle one of these is embedded in a sclerenchyma band which extends between the two surfaces. 3. A. ciliata (fig. 17). The illustration represents the basal portion of an inflorescence-axis bract. Numerous veins are present. These are encased in sclerenchyma sheaths, some of which are continuous with the sclerified adaxial surface. Chlorenchyma is present on the abaxial surface. 92 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN | [vou. 10 4. A. linearifolia. An inflorescence bract is illustrated in figure 23. The abaxial epidermis is sclerified; the mesophyll is divisible into three zones, from outside to inside: chlorenchyma, non-photosynthetiec parenchyma, and _ scle- renchyma. 5. A. macrostachya var. angustior. The outer portion of an inflorescence- axis bract consists of spongy chlorenchyma. The adaxial portion of the mesophyll, in which veins are located, consists of non-photosynthetic parenchyma. A few fibers are present on the phloem poles of larger bundles. 5. A. macrostachya var. macrostachya. The two sections shown (figs. 18, 19) illustrate a great deal of difference between the median region and the most apical portion of an inflorescence-axis bract. At the lower level (fig. 19), the mesophyll contains sclerenchyma on the adaxial face, and a few bands of such tissue on the abaxial face. Chlorenchyma and non-photosynthetie par- enchyma form bands within the bract. At the highest level (fig. 18), the entire mesophyll consists of thin-walled sclerenchyma. 6. A. macrostachya var. robustior (figs. 20, 22)..The inflorescence-axis bract is distinctive in that the ground tissue consists of sclerenchyma, more nearly thin-walled halfway between the surfaces of the bract, which contains pockets of parenchyma, possibly chlorenchyma. In addition to the main series of veins, a few smaller veins are adjacent and exterior to the parenchyma pockets. They have the same collateral orientation as the main veins. The inflorescence bract (fig. 22) is not dissimilar in structure, although it has more thick-walled sclerenchyma, definite chlorenchyma, and bundles adaxial to the chlorenchyma. Orectanthe. An inflorescence-axis bract of Orectanthe sceptrum subsp. occidentalis shows a notable lack of mesophyll differentiation. The entire mesophyll consists of thin-walled parenchyma. Small fibrous bundle caps are present on the phloem poles of the veins. Inflorescence-axis bracts of O. ptaritepwiana showed an identical structure. Achlyphila. The bracts in this plant (fig. 66) are thin, papery inflorescence bracts, not entirely camparable with any of the above because of the distinctive inflores- cenees of this plant. The three bundles are each ensheathed with a fibrous bundle sheath. Discussion. Obvious distinctive modes of bract structure occur in the taxa mentioned above. The lack of mesophyll differentiation in Orectanthe and Achlyphila bracts is intersting, for considerable differentiation occurs in comparable bracts of Abolboda. In. Abolboda, particuiar types of bract structure (depending at least in part on the location of*the bract on the plant and the level of section within a bract) suggests caution in the formulation of differences among the species of Abolboda in bract anatomy. SEPALS With the exception of Achlyphila, all the taxa studied here have keeled sepals. Some Xyridaceae, such as Orectanthe, have both keeled and non-keeled 1960] ANATOMY OF GUAYANA XYRIDACEAE 93 sepals. The non-keeled sepals were found to have approximately the same structure as the winged portion of a keeled sepal (compare, for example, figures 21 and 64), so that separate descriptions are not necessary. The chief variation of significance is the relative distribution and presence of sclerenchyma and (if present) chlorenchyma. To a large extent, the distribution of these tissues in comparison with non-photosynthetic parenchyma cells provides distinctions of taxonomic importance. The patterns are best summarized in terms of species. Abolboda. 1. A. acaulis (fig. 56). The majority of the sepal is composed of lignified parenchyma, in which the three bundles are embedded. Chlorenchyma occurs in the keel of the sepal. 2. A. acicularis (fig. 57). The adaxial face of the sepal is composed of thick-walled sclerenchyma; the pattern of distribution parallels the outline of the bract; the portion adaxial to the keel contains the single vascular bundle. The outer face of the bract contains spherical chlorenchyma cells; a small zone of non-photosynthetic parenchyma is present in the keel. 3. A. americana (fig. 58). The structure of this sepal is virtually the same as that shown for A. acaulis. 4. A. ciliata (fig. 59). The sepal of this species contains sclerenchyma, which is present on the adaxial face and extends toward the keel, where it encloses the midvein. An additional strand of sclerenchyma is present in the most abaxial portion of the keel. Chlorenchyma occurs on the abaxial face of the bract. 5. A. linearifolia (fig. 60). This sepal is distinctive in the limited amount of sclerenchyma present on the adaxial face. The remainder of the bract con- sists of parenchyma cells which are polygonal as seen in transection. The cen- tral portion of this parenchyma contains numerous chloroplasts. Five veins are present in the mesophyll. 6. A. macrostachya var. angustior (fig. 61). This sepal has sclerenchyma distribution similar to that of A. linearifolia, but the adaxial band is wider and is connected with at least a partial abaxial band in the wings (at levels higher than that shown). Sclerenchyma is very thick-walled. Five bundles are present in the non-photosynthetic parenchyma which composes the remainder of the sepal. 7. A. macrostachya var. macrostachya. The sepal in this variety in inter- mediate between those of var. angustior and var. robustior in that the abaxial sclerenchyma band is better developed than in var. angustior. Some of the lateral bundles are embedded in the sclerenchyma of the wings. 8. A. macrostachya var. robustior (fig. 62). The sclerenchyma on the adaxial surface is paralleled (and in places, fused with) a band which parallels the abaxial surface along the wings. The abaxial sclerenchyma extends farther into the keel than in the other varieties. The five bundles are present in the non-photosynthetic parenchyma which comprises the remainder of the sepal. 9. A. sprucei (fig. 63). This sepal shows two distinct zones of sclerenchyma: an adaxial one which parallels the inner face of the sepal, and an abaxial zone in the. keel. Appproximately nine bundles are present. A narrow band of chlorenchyma is present on the abaxial face of the bract. | 94 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Orectanthe. 1. O. ptaritepwiana. The sepal of this species consists wholly of lignified parenchyma, except in the central portion of the keel, where thin-walled parenchyma occurs. This thin-walled parenchyma presumably contains a moderate number of chloroplasts, as it does in O. sceptrum. 2. O. sceptrum. As shown in figure 64, sepals of O. sceptrum subsp. oc- cidentalis are like those of O. ptaritepuiana except that pockets of thin-walled parenchyma, like that in the keel region, occur in the wing regions between veins. Liquid-preserved material was used to demonstrate the presence of chloroplasts in this soft-walled parenchyma. The central portion of a flattened sepal of this species, shown in figure 21, illustrates these same structural features. Achlyphila. The three sepals of Achlyphila disticha (fig. 65) are rounded rather than keeled. They contain about five bundles, which are embedded in the thin-walled parenchyma, composed of cells polygonal in outline, in the central portion of the sepal. This parenchyma contains chloroplasts in moderate numbers. The inner and outer faces of the sepal are sclerenchyma. At lower levels, this sclerenchyma takes the form of hegnified parenchyma which intergrades with the thin-walled parenchyma in the center of the bract, but at higher levels, such as that shown, the sclerenchyma is thick-walled and clearly definable from the chlorenchyma. Discussion. The sepals of Orectanthe are different from those of Abolboda in that (1) they are much larger; (2) they consist wholly of lignified parenchyma cells except for pockets of chlorenchyma in the keel and between some of the veins; and (3) they contain a much greater number of veins, and veins have a scattered distribution in the keel region. Sclerenchyma is present around veins in Orectanthe sepals regardless of location of veins. Sepals of Achlyphila are very simple in structure, corresponding with other seemingly unspecialzed features of this genus. The presence of both an adaxial and an abaxial band of sclerenchyma, together with the lack of a keel, is notable. The species of Abolbod& seem to agree in their tendency toward production of an adaxial sclerenchyma band, and in the relative paucity of sclerenchyma in the abaxial portion of the sepal. Only a single series of bundles is present (unlike the condition in Orectanthe). The patterns of distribution of sclerenchyma, thin- walled parenchyma, and chlorenchyma undoubtedly lend themselves to use as specific criteria, and are self-evident, in the varied patterns illustrated, in Fic. 56-65, sepals, transections taken about midway along length of sepal; half of a keeled sepal is shown for each species. Fic. 56. Aboldoda acaulis, Fic. 57. A. acicularis var. acicularis. Fig. 58. A. americana. Fic. 59. A. ciliata. Fic. 60.. A. linearifolia. Fie. 61. A. macrostachya var. angustior. Fig. 62. A. macrostochya var. robustior. Fic. 63. A. sprucci. Fie. 64. Orectanthe sceptrum. Fic. 65. Achlyphila disticha. Fic. 66. Achlyphila disticha, transection of braet of inflorescence proper. Patterns as follows: fine stippling = thick-walled sclerenchyma; coarser stippling = thin-walled sclerenchyma; cross-hatched = chlorenchyma; white in bundles = phloem; black = xylem; parenchymatous ground tissue left blank. Seale = 2 mm in 1 mm divisions; applies to all figures. 1960] ANATOMY OF GUAYANA XYRIDACEAE v 96 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 sepals of Abolboda. Because of variation in such patterns between the base and apex of a sepal, however, such patterns should be applied with caution. Un- fortunately, no liquid-preserved sepals of Xyris were available for study. The writer suspects that these would show much resemblance to sepals of Abolboda, and would be worth investigating in any case. FLORAL VENATION The flowers of Abolboda and Orectanthe are alike in a number of features. The flowers of Orectanthe differ in having non-keeled sepals, in lacking staminodia, and in having larger parts which are more richly vascularized at least in some instances (corolla lobes, sepals) than the corresponding portions of Abolboda flowers. The general venation pattern in the two genera is suf- ficiently similar so that variations are considered below after a description of a typical venation pattern in Abolboda. Abolboda. The flower of A. linearifolia (figs. 67-81) proved convenient material for a study of venation, and does not appear to differ appreciably from other species in these respects. The levels shown offer a series of selected sections from the base of the flower to the top. Because of venation complexities, a series of sections appeared preferable. The broken line in figures 67-69 is merely used to indicate incomplete separation of a sepal from tissue of the inflorescence receptacle at those levels. At the lower level indicated (fig. 67) a vascular cylinder can be seen. The bundles of the cylinder have branched, at a level lower than that shown, to supply the approximately five traces present in each sepal. At the next level (fig. 68) sepal traces are in their characteristic positions, and divergence of median corolla bundles is indicated. Only ovary bundles are present on- the central area a short distance above this level. Five traces depart to each third of the corolla tube; these thirds correspond to the three corolla lobes at a higher level. This is more evident in figure 69, which shows two of the three staminodia that will be alternate with these thirds of the corolla tube. There is a clearly definable midvein in each third of the corolla tube (see particularly figure 72). At the level shown in figure 69, as well as in figure 70, the nature of ovary yenation is evident. There are three dorsal carpellary traces and three ventral ones, although the latter are fused together at these levels. At the next higher level (fig. 71) separation of the axial portion of the ovary into three placentas, each with a vein (or in some preparations, a pair of veins) is shown. Fias. 67-81. Abolboda linearifolia, successive transections of a flower. Sepals are omitted in figures 72—75, and style only is shown in figures 76-81. Fic. 67. Departure of traces to sepals. Fic. 68. Departure of traces to corolla. Fic. 69. Base of ovary. Fie. 70. Level at which placentas are united; note three staminodes, each opposite a locule of ovary. FIG. 71. Level at which placentas are separate. Fic. 72. Upper end of ovary locules. Fic. 73. Level of ovary crests; note the triangular style base in center, the three staminodes just inside the corolla, and the two style appendages. Fic. 74. Departure of stamen traces; base of anthers; note several traces in each style appendage. Fie. 75. Level of stamen connectives; note cylindrical conformation of bundles in style of appendages. Fic. 76. Departure of style ap- pendages from style. Fias. 77-80. Branching of bundles in upper portions of style. Fic. 81. Stigma. Vascular bundles in all figures represented by circles (or modified shapes). Seale = 2mm in 1 mm divisions. d7 ANATOMY OF GUAYANA XYRIDACEAE 1960] 98 MEMOIRS ‘OF THE NEW YORK BOTANICAL GARDEN |vou. 10 Figure 72 shows the termination of the ventral carpellary traces (below the level shown). The dorsal traces turn inward as the ovary narrows into the style, and only the three dorsal traces are present in the style base (fig. 73). In figure 73, the ovary appears fragmented into three portions (other than the style and style-appendages), and the larger triangular shapes represent the ovary crests. At the next level (fig. 74) departure of stamen traces, by branching from the median corolla traces, can be seen. The uppermost of the staminodia shown terminates at this level. In figure 75, the three corolla lobes become free from each other. Concomitantly with their greater widening into an imbricate conformation, there is considerable ramification of veins. The upper tips of two of the ovary crests may be seen at this level. The following figures show the changes in vascularization of the style. In figure 76, departure of ovary appendages may be seen. These appendages are recurved, so that description of changes in their anatomy from departure to tip preceeds in the reverse order, in terms of figure numbers. The bundles that depart into the ovary appendages are branches of the three style traces, not the traces themselves. One of the appendages is very short; the other two (fig. 75) show an alteration in vascularization. A cylindrical organization of the bundles is attained. At a lower level (fig. 74), these cylinders subdivide into several bundles, and at the tip of the appendages (fig. 73) only a single bundle is present. This bundle, in each, does not extend all the way to the appendage apex. Returning to the style proper (fig. 77), one may see that the three veins branch. This branching continues (figs. 78-81) so that numerous vein-endings enter the ultimate fringes of the stigma. Development of a central cavity in the style is illustrated in figure 80. Variation in venation among the species of Abolboda seems to occur only in the number of bundles per sepal (see above) and the relatively abundant or few veins in the corolla, depending largely on corolla size. The question naturally arises, what interpretation of the ovary appendages seems most appropriate in terms of their venation, position, and _ histology. They cannot be regarded as equivalent to two of the three stigmas (if two are present), on account of their venation. They represent, rather, branches of the style bundles. The vein in each appendage does not terminate as a hydathode, nor do the contents of cells in the appendage suggest a nectary function, al- though this possibility should not be ruled out. The function of these structures —if they do have any prominent function—may well be related to a pollination mechanism in some way, but without further knowledge, speculations concerning function would be pointless. A feature of some interest is the nature of bundles in the very large ovary appendages of Orectanthe. In O. sceptrum, the bundle was observed to be large and rather diffuse, with isolated patches of xylem and phloem cells. . In respects other than numbers of veins and absence of staminodia, the resemblance between Abolboda and Orectanthe flowers are close, and ovary and_ style of Orectanthe flowers have a venation identical to that in Abolboda. Figs. 82-87. Achlyphila disticha, successive transections from flower. Sepals omitted in | figure 86, and only style base shown in figure 87. Fic. 82. Pedicel. Stippling represents thick- walled sclerenchyma. Fic. 83. Departure of sepal traces. Fic. 84. Base of petals. Fie. 85. Departure of stamen traces. Fic. 86. Ovary at level of ovules. Fic. 87. Style base. Vascular bundles in all figures represented by circles (or modified shapes). Scale = 2 mm in 1 mm divisions, 1960 | ANATOMY OF GUAYANA XYRIDACEAE ao ‘ - 100 MEMOIRS,OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Achlyphila. The venation of the flower of Achlyphila is relatively simple in comparison with that of Abolboda. Fewer levels therefore suffice to show the principal features, and are shown in figures 82-87. The pedicel, illustrated in figure 82, contains two circles of bundles. The outermost of these circles departs into the bases of the three sepals (fig. 83), in such a way that five traces are present in each sepal (fig. 84). Successive levels (figs. 84, 85) illustrate that above the level of sepal departure, the three whorls—petals, stamens, and gynoecium— become separate at approximately the same point. As in Abolboda, the stamen traces branch from the vein which can be demarcated as the midvein of each petal. No traces indicate any vestige of three other stamens, or of staminodia, which might be assumed to have been present primitively. About five traces are present in each petal, although these probably originate from about three traces in each petal base. The venation of the ovary is identical to that of Abolboda, as a higher level (fig. 86) indicates. As in Abolboda, widening of the petals is concomitant with branching of veins. Ridges, which fit into interstices between the sepals, were observed on the abaxial surface of the petals, and presence of these ridges alters the appearance of petal vascularization somewhat, for these ridges are vascularized. The venation of the style is as in Abolboda, except that no branching of the three veins takes place, as no appendages are present. The similarities between Abolboda (e.g., A. linearifolia) and Achlyphila in respect to floral venation are quite striking. The only major differences are in the relative levels at which veins depart. Staminodia and ovary appendages are absent in Achlyphila, and three, rather than two, sepals are present (although some species of Abolboda also have three sepals). FLORAL HISTOLOGY Certain details of floral anatomy other than venation are worthy of descrip- tion here because of their importance in suggesting specific and generic relation- ships. Some of these have been mentioned in connection wth sepal structure. Corolla Lobe. Corolla lobes, as might be expected, are thicker in their central portion than at their margins. The maximum width of corolla lobes was found to offer a convenient point of comparison. The corolla lobe of Abolboda linearifolia (fig. 92), like those of the other species of Abolboda, has a maximum width of ap- proximately four mesophyll cells. The maximum mesophyll width in corolla lobes of Orectanthe sceptrum subsp. occidentalis (fig. 91) proved to be about 10 cell layers; these cells are much larger than those of Abolboda. In Achlyphila disticha, the maximum width (excluding ridges) appeared to be about 8 layers, - as shown in figure 938, with a lgnified sheath around veins. Fias. 88-90. Portions of transections of anthers. FiG. 88. Abolboda linearifolia. Fa. 89. Orectanthe sceptrum. Fia. 90. Achlyphila disticha. Fies. 91-93. Portions of transections of corolla lobes or petals. Fic. 91. Orectanthe sceptrum. Fie. 92. Abolboda linearifolia. Fig. 93. Achlyphila disticha. Seale for figures 91-93 is shown beside figure 92. All seales are in 1 mm divisions. 101 a peste, > 3 oe po) Ml) Qs age / ate Beedqensey, (/ = Z oe BE “ @ g 6 ss S20 oe? ANATOMY OF GUAYANA XYRIDACEAE 1960] AS ar ) a c Micappy ye > b Sz RX LY rr 5 m4 ‘ ) .) hy TH ee a cx fs YY G 89 4 J S) aly Wr, Dip > M ( wal NN ZL \, 88 G7; Uff : RCUTMinee G) @ S b Zant Sta S 2 = —@\ Fig. 101. Abolboda americana, embryo from sagittal sections of embryo sae. Note basal and hypobasal cell and haustorial ‘‘flaps.’’ Endosperm shows many nuclei (stippled) per cell, starch grains (with shrunken hilum indicated), and protein storage globules (black). Micro- pyle below. Fic. 102. Orectanthe sceptrum. Embryo, same view as that shown in figure 101. Fig. 103. O. sceptrum, section of seed coat from longitudinal section of seed. Section repre- sents a portion between the prominent wings on the seed, which are therefore not involved in this section. Fie. 104: Abolboda macrostachya var. angustior, transection of capsule valve. 1960 | ANATOMY OF GUAYANA XYRIDACEAE 107 showed that the outer thin-walled layers, without contents, are derived from the outer integument, whereas the inner pair of cells with occluded lumina are derived from the inner integument. From very casual observations on seeds of Xyris, the writer is of the opinion that the respective layers in Yyris actually have a similar origin, and that the interpretation of Weinzieher is open to question. The differences in seed structure between Abolboda and Orectanthe appear to be very distinctive expressions of the same basic pattern, and may be used both to separate the genera and to emphasize an underlying similarity. The writer believes that seeds of Yyris, upon further study, will probably also reveal similar conspicuous variations on a pattern common to the whole family. Study on seeds of Achlyphila is needed for the same reasons. Endosperm. Weinzieher (1913) and Netolitzky (1926) indicate rounded compound starch grains and protein spheroids in endosperm of Xyris. Presumably both of these accounts are based on Weinzieher’s study of X. indica. All the species for which material was available of Abolboda (fig. 101) and Orectanthe showed these features. Immature stages in the endosperm of A. sprucei showed the multinucleate condition of endosperm cells very clearly. Embryo. Weinzieher (1913) has illustrated stages in the development of the embryo of Xyris indica. The embryo consists of relatively few cells. A broad, flat cotyledon is evident, but Weinzieher figures no differentiation of a suspensor or any meri- stematic region. Features like these occur in Abolboda, as illustrated for A. americana in figure 101. A basal and a hypobasal cell are evident, however. The cotyledon consists of large cells, with dark-staining contents that suggest the haustorial function of this structure which is closely appressed to the endosperm. At the edges of the cotyledon, flap-like extensions are evident. The embryo of Abolboda americana shows no differentiation of an apical area, and is larger and composed of more numerous cells than that of X. indica. Significantly, the embryo of Orectanthe sceptrum (fig. 102) shows precisely the same features as the embryo of Abolboda americana, but is larger, corresponding to the larger size of all organs in this species. A basal and a hypobasal cell appear to be present. If a meristematic area is present, it is not appreciably differentiated from other areas of the embryo. Unfortunately, no embryos of Achlyphila were available. Likewise, no stages in gametogenesis or early embryo development were visible on account of the nature of preservation. POLLEN Methods Pollen grains of Yyridaceae are subject to decomposition by various reagents. As Erdtman (1952) notes, his acetolysis method degrades the exine. Sodium Fic. 105. Orectanthe sceptrum, transection of capsule valve. Seale beneath figure 104 applies to figure 105; figure 102, 103 at same seale (between those two figures). All seales in divisions of 0.1 mm. 108 MEMOIRS, OF THE NEW YORK BOTANICAL GARDEN [von. 10 hydroxide, a reagent commonly used in pollen-preparation techniques, partly or wholly dissolves the exine. Except for two specimens (figs. 112, 120) in which NaOH treatment was very gentle and merely revealed some textural differen- tiation in the exine—probably because of preferential solubility of certain parts—this substance proved useless. Even the pollen of fixed material which formed the main basis for this study was, in some eases, altered. Because of this extreme sensitivity to a wide variety of reagents, the simplest methods for preparation of pollen grains for observation appeared to be (1) sectioning and staining of flowers (with their included pollen) according to the techniques described at the beginning of this paper; and (2) transferring fixed anthers through an alcohol series to absolute ethyl alcohol, staining in safranin dissolved in absolute ethyl alcohol, and following this by transferring anthers to xylene and making whole mounts of grains in Canada balsam. Undoubtedly both of these techniques resulted in dehydration and perhaps other changes which could alter size and possibly induce artifacts. However, a number of grains in each of the collections studied appeared to be reasonably unaffected, and were con- sidered suitable for study. Note should be made of the fact that the spines (or other excrescences of the exine) that stain bright red are seemingly unchanged by any of the reagents mentioned, and data concerning these are without doubt quite reliable. The exine wall, which stains green with a safranin—fast green combination, is the structure that is sensitive to reagents. Because the grains are large, and few in number per anther, because good material (grains from anthers just before or at anthesis) were necessarily limited, and because many grains were collapsed, or degenerate, the number of grains that could provide reliable measurements was extremely small. The data on pollen grain diameter below represent, therefore, averages of measurements of only those grains that the writer judged to be turgid and otherwise unaltered in form. On account of the exceptional susceptibility of pollen grains in this family to various changes, truly exact data can probably be obtained only from fresh material. Such material is virtually impossible to view because of the remote locations in which these species grow. Terminology for pollen morphology used below follows the usage of Erdtman (1952). Pollen Grain Size. Pollen grains of Abolboda, Orectanthe, and Achlyphila are sphaeroidal, a fact which may be related to their nonaperturate condition. Because of this shape, only one dimension is necessary. Dimensions do not include the spines or other excrescences. Species Diameter in microns Abolboda acaulis (fig. 106) 84 A. acicularis var. acicularis (fig. 107) 96 A. bella (fig. 108) : 10 A. ciliata (fig. 109) 96 A. ebracteata (fig..110) 90 A. grandis var. guayanensis (fig. 111) 150 A. linearifolia (fig. 112) 120 A. macrostachya var. angustior (fig. 118) 1595 A. macrostachya var. macrostachya (fig. 119) 105 1960] ANATOMY OF GUAYANA XYRIDACEAE 109 A. macrostachya var, robustior (fig. 120) ) 180 A. sprucei (figs. 114-117) 120 Orectanthe ptaritepwiana (figs. 121-124) Steyermark & Wurdack 523 165 Steyermark & Wurdack 1221 165 Maguire et al. 32823 140 O. sceptrum (figs. 125-127) 140 Achlyphila disticha (figs. 128, 129) a2 Within the genus Abolboda, some taxa seem obviously to have larger pollen grains than others; species with exceptionally large grains include A. linearifolia, A. macrostachya (at least in part), A. grandis, and A. sprucet. The pollen grains of Orectanthe are markedly larger, in general, than those of Abolboda. The markedly smaller diameter of Achlyphila pollen grains is interesting; this size is comparable to the sizes reported by Erdtman (1952) for species of Xyris. Pollen grains of Xyris, as described below, do differ in morphology from those of Achlyphila. Previous reports on pollen-grain diameter include those of Malme (1933) on ‘‘Abolboda’’ (now Orectanthe) sceptrum (‘‘about 105 »’’) and Erdtman (1952) for Abolboda poarchon (‘‘105 p’’), A. pulchella (‘'105 w’’), and A. vagi- nata (‘‘at least 75 p’’). Maguire, Wurdack et al. (1958) give the following measurements: Abolboda bella, 120-140 »; A. paniculata, 140-150 »; and Orec- tanthe (as a genus), 160-250 up. Pollen Grain Wall (‘‘sporoderm’’). Sectioned_ material showed two distinct parts of the pollen grain wall in all taxa: the inner portion, which stains brightly with fast green, and the spines, or Other ornamentation (including the minute pila on the surface of the inter- spinal areas), which stain bright red with safranin. These two portions appear to correspond with Erdtman’s definitions of nexine and sexine respectively, except that a thin layer, which stained pale green, on the inner surface of the pollen grain wall in Abolboda could be interpreted as the intine. This interpreta- tion was suggested by observation of germinating pollen grains on a style of A. grandis. In these, this layer covered the emerging pollen tube. Such a layer could not be distinguished in pollen grains of Orectanthe, possibly because of difficulties in preservation, but a comparable layer did appear present in grains of Achlyphila (fig. 128). The nexine in sectioned material and in whole mounts appears to stain homogenously in most preparations. In two preparations in which sodium hydroxide had been employed (fig. 112, 120), radial fibrillae or striae appeared to be present in the outer two-thirds of the nexine. This appearance may have been caused by differential solution of materials in the nexine, or by expansion of the wall by means of the treatment. Among the various taxa, the nexine thickness appears to be.a distinctive character. In most of the Abolboda species studied, it appears to range between 5 and 7 uw. It is much thicker than that in A. grandis var. guayanensis, A. macrostachya var. macrostachya, A. macro- tachya var. robustior, and A. sprucei. Certainly the great thickness of the nexine of A. grandis var. guayanensis and A. sprucei represents a distinctive character- istic. In Orectanthe, the nexine does not appear to exceed 5 pw in thickness. In Achlyphila, the width of this layer is about 3 x. 110 MEMOIRS, OF THE NEW YORK BOTANICAL GARDEN [vou. 10 » dae e &*7 ogap? Ts Figs. 106-117. View of pollen grains in species of Abolboda. All figures except figures 113, 115-117 represent a portion of a median microtome section of a pollen grain, and have been drawn to show one of the large ornaments; intine is indicated below by sparser stippling. Fic. 106. A. acaulis.' Fic. 107. A. acicularis var. acicularis. Fic, 108. A. bella. Fie. 109. 1960 | ANATOMY OF GUAYANA’ XYRIDACEAE 111 The outermost layer of the pollen grain wall, or sexine, is basically a thin membranous layer which may be partly detached in damaged grains. Minute pila (seen in surface view, figs. 113, 129) are embedded in this membrane. A feature that seems quite important to the writer does not seem to have been mentioned in the literature on Xyridaceae pollen grains, namely, that these minute pila are, in fact, homologous with the large spines or knob-like ornaments in Abolboda and Orectanthe. These larger excrescences are deposited on the membranous layer just as the pila are, and appear to be composed of the same material, despite the vast difference in size between the two types of ornamenta- tion on the same grain. Rarely, as in Orectanthe sceptrum (fig. 125), knobs transitional in size between these two categories may be observed. The pila in Orectanthe are noticeably larger than those in Abolboda. Pollen grains of Achlyphila (figs. 128, 129) show the largest pila of any of the grains studied here. In Achlyphila, the pila tend to be aggregated: in flake-like patches. In places between these patches, the pilate layer is absent. In Abolboda, a thin hyaline layer was observed immediately beneath the pilate layer. Such a layer may also occur in pollen grains of Orectanthe and Achlyphila. The spines on pollen grains of Abolboda are so characteristic of that genus, and are so subject to distinctive variations in the species, that they have been figured for all the species studied (figs. 106-120). Basically, as in A. acaulis or A. linearifolia, they have an inverted funnelform shape. Sodium hydroxide treatment (figs. 112, 120) reveals that lacunae, not visible in other preparations, are probably present in the spine base. Some species show markedly narrow spines: A. bella (fig. 108), A. acicularis (fig. 107), and A. ciliata (fig. 109). Very short, wide spines occur in A. ebracteata (fig. 110) and A. macrostachya var. robustior (fig. 120). The three varieties of A. macrostachya appear different on the basis of spine size and shape. Within a single flower of A. spruced, various alterations in spine shape were noted. The basic condition seems to be that shown in figure 114, but multiple spines on a single base (fig. 115), very much reduced lobes (fig. 116), and merely the vestige of the spine-base (fig. 117) were also observed. Erdtman (1952) has illustrated the spines of pollen grains of three species and given dimensions:A. poarchon (spines 8.3 m long; basal diameter 5.5 p), A. pulchella (spines 9.3 » long; basal diameter 8.3 »), and A. vaginata (spines 8.3 » long; basal diameter 8 »). Erdtman figures spines in the latter species which appear about like the spines in A. macrostachya var. angustior. The comparable ornamentation on pollen grains of Orectanthe is generically different from that of Abolboda. In Orectanthe, the emergences take the form of a large knob, or other shape much wider and more blunt than the spines of Abolboda, with the possible exception of the much smaller structures in A. sprucet. In O.- ptaritepuiana (figs. 121-124) a variety. of shapes was noted, varying with the collection from which the grain was taken: short, with a rounded apex (fig. 121), elongate (fig. 122), with a curved apex (fig. 123), or with folds or bulges on the sides (fig. 124). Evidently considerable variation is ep A. ciliata. Fig. 110. A. ebracteata. Fic. 111. A. grandis var. guayanensis. Fig. 112. A. linearifolia (from NaOH-treated specimen). Fic. 113. Same, surface view of interspinal sexine ornamentation. Fic. 114. A. sprucei. Fig. 115-117. Same, variations (within a sin- gle flower) or ornament shape. Scale = 5 y. ' 112 MEMOIRS ,OF THE NEW YORK BOTANICAL GARDEN [von. 10 , Io? prsrra-Fr tle = Pes AT ‘ »-. = Lo Pe. oN wo F Fie rear: SIS" SY OAS, XO ES $y? Bee =a) a ot O Nee Ue: Si te Ea) 3 “ae WERE = ye Pay atte ; H Sy aed “e i) 26 brace: OS aS? D wet ae 9, B 3 YS ri \2S sie wee MS Fics. 118-129. Views of pollen grains. Conventions as in figures 106-117. Fic. 118. Abolboda macrostachya var. angustior. Fie. 119. A macrostachya var. macrostachya. Fie. 120. A. macrostachya var. robustior (from NaOH-treated specimen). Fie. 121. Orectanthe ptaritepuiana, Fics. 122-124. The same, variations in ornament shape. Fie. 125. O. scep- 1960 | ANATOMY OF GUAYANA XYRIDACEAE PS present in these characters. In preparation of pollen grains in this species, the knobs appeared to be sunken in slight depressions; this may well be an artifact of fixation or preparation. The knobs of Orectanthe sceptrum pollen grains are different from those of O. ptaritepwiana in that the apical portion is widened, so that a capstan-like shape is achieved. The only variation in structure of these knobs is found in the formation of additional small knobs on the base (fig. 127) as compared to the normal condition (fig. 125), or the formation of a roughened base (fig. 126). Discussion. In summary, the pollen grains of the three genera show excellent generic characters. The presence of large excrescences in Orectanthe relates this genus closely to Abolboda, but the differing shape is a good generic character. Like- wise, the larger pila in the surface between emergences differentiates Orectanthe from Abolboda. The absence of such large emergences in Achlyphila, and the grouping of pila into flake-like aggregations on the exine surface, marks this genus off from other genera of Xyridaceae. Within Abolboda, differing exine widths and distinctive spine shapes and sizes serve to differentiate the species, so that of the species studied, most could be identified by means of the pollen grains alone. Similar considerations apply to distinctions between the two species of Orectanthe. Erdtman (1952) has emphasized the great distinction between Abolboda and Xyris in pollen-grain structure. Much of this distinction may be attributed to the large spines on the surface of Abolboda pollen grains. The lack of such larger ornaments on grains of Achlyphila, which are (like those of Abolboda, but unlike those of Xyris) nonaperturate, may vitiate this contrast, and pro- vide an intermediate form. The pilate layer in pollen grains of Abolboda, Orec- tanthe, and Achlyphila probably has a counterpart in Yyris. The presence of the pilate layer, although different in each genus, tends to provide a common characteristic, as does the rather thick nexine, lacking in hpophilic compounds. The writer suspects from Erdtman’s reports of both ‘‘OL’’ and ‘‘LO”’ sexine patterns that a layer of united pila may compose the sexine in at least some Xyris pollen grains. The exine of Achlyphila seems midway between that of Abolboda and Orectanthe on one hand, and Xyris on the other. The relatively thick, baculate sexine and the thin nexine of Xyris pollen grains do not seem at odds with the condition in Achlyphila. Probably the three major patterns represent striking variations on the same basic pattern. Careful work is needed both to provide additional data on Xyris and to demonstrate whether the affinity Erdtman (1952) suggests between the grains of the Abolboda-type and those of Liliaceae and Zingiberaceae is close or not. SUMMARY The writer has found that an attempt to summarize anatomical character- istics of genera of Xyridaceae in chart form is virtually impossible because (1) trum. ‘Fics. 126-127. The same, variations in ornament shape. Fie. 128. Achlyphila disticha. Fic. 129. The same, surface view of sexine ornamentation (pila). Seale = 5 w#. \ 114 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 there are exceptional species within a genus; (2) reference to particular con- ditions in lengthy anatomical terms, or with the aid of illustrations is necessary ; and (3) some relatively subtie characteristics cannot be summarized in this man- ner. The most important anatomical lines of evidence are cited below, and the reader is referred to the foregoing descriptions, particularly to the ‘‘discussion”’ section terminating each of the major portions above. Specific Characteristics. 1. Abolboda. Characteristics of leaf anatomy and stem anatomy suggested recognition of two species-groups, termed here ‘‘smaller-stemmed’’ abolbodas and ‘‘larger-stemmed’’ abolbodas. Recognition of these as subgenera would probably be premature. Within each of these groupings, leaf and stem anatomy suggest specific characteristics also. Characteristics other than these do not emphasize the two species-groups as much as they suggest characteristics of individual species. The large stele of roots defines A. macrostachya, but genuine- ly remarkable specific characteristics are offered by the peculiar patterns of endodermis-cell thickening. In the inflorescence axis, thickness of sclerenchyma ring, presence of fibrous sheaths on pith bundles, and sclerification of epidermal cells offer a number of characteristics. Distinctive patterns in number of bundles, size, sclerenchyma and chlorenchyma presence and distribution offer further characteristics in sepals, and, to a lesser extent, in bracts. Perhaps the best specific characteristics are evident in pollen in such respects as shape and size of spines, nexine thickness, and diameter. 2. Orectanthe. Anatomical features by which O. ptaritepwana may be differentiated from O. sceptrwm include relative lignification of the central portion of the stem; thickness, epidermal sclerification and shape of bundles in leaves; distribution of parenchyma in the sepals; size and sexine ornamentation of pollen grains. Generic Differences. The closest relationships among genera of Xyridaceae is that between Abol- boda and Orectanthe. Segregation of the latter genus is well deserved, however. Orectanthe has larger stems with a distribution of sclerenchyma different from that in Abolboda. Orectanthe has more bundles in its leaves, more bundle-sheath extensions, a tendency towards abaxial hypodermis production, and no thorn- hike leaf-tip with peculiar tracheids. It has spongy chlorenchyma and lignified bundle sheaths in the cortex of the inflorescence axis. It has distinctive sepal size and anatomy, and the inflorescence-axis bract is notably lacking in scleren- echyma or other differentiation. Orectanthe lacks staminodia, has more elaborate corolla venation, and a massive anther connective. Capsule-valve anatomy and especially seed anatomy provide excellent characteristics to distinguish the two genera. Pollen morphology provides an exemplary instance oni a generic degree of distinction between Abolboda and Orectanthe. Achlyphila could probably be defined, on the basis of present knowledge, on a number of anatomical features. The root offers peculiar endodermis characters and vessels well toward the interior of the core, which, however, lacks central vessels. The stem shows modifications corresponding with its rhizomatous habit, such as a stem endodermis. The leaves have a peculiar equitant habit, and the 1960 | ANATOMY OF GUAYANA XYRIDACEAE 115 central core of lignified parenchyma in upper portions is peculiar. The flattened inflorescence axis, lacking veins in the ‘‘pith’’ region, is distinctive. Sepal anatomy could be used to separate Achlyphila from the other genera, and floral venation reveals both a lack of staminodia and a choripetalous condition. Pollen is peculiar because it combines a peculiar patch-like presence of long pila with a lack of major ornaments and a nonaperturate condition. Detailed comments cannot be made on Xyris, because it has been excluded from this study. Such characteristics as lack of ‘‘pith’’ bundles in the inflores- cence axis, distinctive seed anatomy, and the peculiar nature of the monosulcate pollen provide features which, as extended descriptions would show, amply enforce the generic distinction of Xyris within the family. Composition of Xyridaceae: Relationships of the Four Genera. In respect to either gross morphology or anatomy, three relatively isolated groups are apparent: (1) Abolboda and Orectanthe; (2) Achlyphila; and (3) Xyris. Before showing the anatomical common denominators among these groups, similarities among pairs of genera may be mentioned. Abolboda and Orectanthe, as mentioned above, are amply distinct, but the resemblances are close. For example, the presence of style appendages and their venation pattern is a striking similarity. Winged seeds occur both in Orectanthe and, to some degree, in Abolboda macrostachya. Seed-coat anatomy of the two genera reveals two variations on the same basic pattern, as do the embryos. The inflorescence axis in both genera contains bundles in the ‘‘pith’’ region. Most significantly, perhaps, both genera possess pollen grains in which an outer layer contains both minute pila and large spine- or knob-like ornaments. Abolboda.resembles Xyris in exhibiting a similar range of anatomical char- acteristics in roots. Equitant leaves and compound bundles in leaves occur in both genera. The embryo of Abolboda, although larger than that of Xyris, is very similar. Better knowledge of seed anatomy in Xyris might reveal similar- ities in this respect. Abolboda is like Achlyphila in relatively few aspects, because the relation- ship of Achlyphila to Xyris appears closer. In addition to features common to all genera mentioned below, one may mention resemblances in epidermal relief in flowers, identical ovules, similarities in floral venation, and certain features of root anatomy and inflorescence-axis structure. Relationships between Achlyphila and Xyris include those of equitant leaves and their anatomical consequences, the presence of sclerified epidermis and undulate epidermal contours on leaves, the lack of bundles in the ‘‘pith”’ region of the inflorescence axis, and resemblances in pollen. The pollen of both genera lacks the differentiation of sexine into major and minor ornaments, and is relatively small in diameter. Review of anatomical characteristics reveals a number in which the genera are alike, or in which the genera appear as variations on a common basis. One significant feature is the presence of vessel elements with simple perforation plates in all parts of the plant. As the work of Cheadle (1942) shows, this high degree of specialization occurs in only a few families of monocots. The amount of similarity among the three genera in this study in respect to floral venation is remarkable, and study of Xyris might enable inclusion of that genus in this respect. The choripetalous condition in Achlyphila is really not % 116 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 appreciably different from the tubular condition of the corolla in Abolboda and Orectanthe, and the venation reveals no appreciable alterations in this respect. To be sure, Achlyphila is less specialized in its choripetalous condition, but one might assume that presence of staminodia was a primitive character in Abolboda. Certainly the development of style (‘‘ovary’’) appendages in Abolboda and Orectanthe are advanced characteristics, whatever their signifi- cance may be. Unfortunately comparable data is not available on seed anatomy for the four genera, but I suspect that this will reveal a common basis upon which variations have been evolved. Similar considerations also apply to morphology of the embryo; the three genera for which this is now known all have the same peculiar type, with variations chiefly in size. The most significant anatomical character in Xyridaceae with respect to difficulties in interpreta- tion might seem to be pollen morphology. Much has been made of the difference between Abolboda (with Orectanthe) and Xyris in these respects. Because monosuleate pollen is so basic in the monocots, the nonaperturate condition in Abolboda, Orectanthe, and Achlyphila would seem to be derived. If this were true, one might also expect a more primitive exine condition in Xyris also. The lack of differentiation of the sexine elements, and their even distribution, appear to be unspecialized. Exine in Achlyphila consists of only one type of sexine element, small pila arranged in flake-like patterns. This ornamentation is close to that of Yyris, in the writer’s opinion, although Achlyphila is like the other two genera in the nonaperturate condition. The most logical explanation of the exine in Abolboda and Orectanthe seems to be that differentiation into two types of elements—minute pila and large spines or knowbs—has taken place; this differentiation most likely would have started from a condition like that found in Achlyphila. The relatively thick nexine, lacking in lipophile com- pounds, offers a common basis among the genera. More study of Xyris pollen is required before a definite phylesis can be suggested, but the above inter- pretations appear most likely at the present moment. The reader will note that the taxonomic conclusions expressed above bear a very close resemblance to those of Maguire, Wurdack, et al. (1958, 1960). The writer agrees with these authors in their conclusion that the four genera of Nyridaceae are highly distinctive, but show more common resemblance than any resemblance of any one to a genus of another family. The writer was pleased ts find, in addition, that his anatomical material lends such clear-cut support to specific and even subspecifie (Abolboda macrostachya) designations of these authors. Literature Cited Arber, Agnes. 1925. Monocotyledons. 1-258. Carlquist, S. 1958. Anatomy of Guayana Mutisieae. Part II. Mem. N. Y. Bot. Gard. 10: 157-184. Cheadle, V. I. The occurrence and, types of vessels in the various organs of the plant in monocotyledons. Am. Jour. Bot. 29: 441-450. Erdtman, G. 1952. Pollen morphology and plant taxonomy. 1-539. Maguire, B., Wurdack, J. J. & collaborators. 1958. The botany on the Guayana Highland— Part ILD... Mem. N.-Y.-Bot. Gard 10:" 1-156. 1960. The botany of the Guayana Highland—Part IV. Mem. N. Y. Bot. Gard. 10: 000-000. Pages to be supplied in proof. 1960] ANATOMY OF GUAYANA XYRIDACEAE TET Malme, G. 1925. Xyridologische Beitrage. Ark. Bot. 19(13): 1-8. . 1933. Beitrage zur Kenntniss der sidamerikanischen Xyridazeen. Ark. Bot. 25(12): 1-18. Netolitzky, F. 1926. Anatomie der Angiospermensamen. Jn: K. Linsbauer [ed.], Handbuch der Pflanzenanatomie II. IOD: 1-364. Solereder, H. & Meyer, F. J. 1929. Xyridaceae. In: F. J. Meyer [ed.], Systematische Anatomie der Monokotyledonen 4: 36—50. Weinzieher, S. 1913. Beitrage zur Entwickelungsgeschicte von Xyris indica L. Flora 106: 393-482. ANNOUNCEMENT Beginning 1 January 1960, all back issues of Mremorrs OF THE NEW YorRK BOTANICAL GARDEN will be handled exclusively by Stechert-Hafner, 31 East 10th Street, New York 3, N. Y. Current issues will be han- dled as heretofore by the Garden. Claims for missing issues should be directed to the Garden. Stechert-Hafner will perform the same service for back issues of all serials pub- lished by The New York Botanical Garden and will reprint parts that have become exhausted. Prices of parts and volumes may be had on application to Stechert- Hafner. a ane y Paras ae A a ss as > ‘ i ‘ “4 Diat Gr me ‘ . Pl te on * he ; > aks ty ble re cn ey TS a J : i. . ~ a ot eed 5 r . : | ; : . } : | é, | VotumME 10, Numser 3 EW YORK BOTANICAL GARDEN | - « on of Spathiphyllum (Araceae)... Grorce S$. Buntinc 1 * 48 N 7 vt i The Memoirs of The New York Botanical Garden are issued at irregular intervals in parts of various sizes. Approximately 500 pages will complete a volume. The subscription price of volume 10 is $10.00. Number 3 may be purchased separately for $2.00. Authors of papers may obtain separate copies of their contributions, printed at the same time as the issue, at cost price. For further information address the editor: H. W. Ricxetr The New York Botanical Garden New York 58, N. Y. NOV 1960 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN 10 (3) :1-53 A REVISION OF SPATHIPHYLLUM (ARACEAE) GEORGE S. BUNTING Missouri Botanical Garden St. Louis, Missouri INTRODUCTION The genus Spathiphyllum forms an inconspicuous element in the flora of tropical America, although it is widespread from Mexico to Peru and Brazil. Two species occur only on Pacific islands, one on Cocos Island (some 300 miles off the Costa Rican coast) and another in Indonesia and the Philippines. Most species of Spathiphyllum grow at elevations between 100 and 1000 m above sea level, frequently in the foothills of mountains or in river valleys. They are plants of permanently wet or moist, shady places that may be inundated at times. They often form colonies along the margins of rivers and streams. Attempts to identify recent Guayanan collections and other material of Spathiphyllum suggested the need to further investigate the classification of this genus. Subsequently, about 1000 specimens were critically examined. Deep ap- preciation is expressed to the directors and curators of the many herbaria from which specimens were borrowed.! HISTORY OF THE GENUS Before the publication of Spathiphyllum in 1832, two species had been de- seribed that are now included in this genus. Jacquin in 1790 (Collectanea 4: 118) described Dracontium lanceaefolium. In 1803, Pothos cannaefolia was clearly illustrated and characterized in Curtis’s Botanical Magazine (pl. 603). Schott, having described Spathiphyllum in 1832 (Melet. Bot. 22), listed ‘‘Sp. laneeaefolium Schtt. (Dracontium lanceaefolium Jacq.)’’ and ‘‘Sp. sagittae- folium Schtt.’’ It appears that he followed Sprengel, who in his revision of Linnaeus’ Systema naturae (Syst. Veg. 3: 766. 1826) had included these two species in the genus Dracontium along with D. podophyllum L. Schott correctly excluded the last species. The two species included with the original description of Spathiphyllum are not congeneric, though both have trimerous flowers with perianth, a superficially acaulescent habit, and long petioles geniculate at the apex. Schott’s treatment was followed by Endlicher (Gen. Pl. 1: 240. 1836) and by Kunth (Enum. PI. 3: 83. 1841). . In 1849, Liebmann described the genus Hydnostachyon (Vid. Meddel. 1-2: 23), differentiated from Spathiphyllum by its unilocular ovary. Critical examina- tion of the specimens referred to that genus fails to verify this unilocular condi- tion. Schott placed Hydnostachyon in synonymy under Spathiphyllum in 1853. 1Abbreviations for herbaria used in the text are those of Lanjouw and Stafleu, Index Herb. 2(1):106-117 (1952). bo MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 In 1852, Carl Koch, impressed by the truncate, cuplike perianth of a plant then cultivated as ‘‘Pothos cannaefolia’’ and ‘‘Monstera cannaefolia,’’ desig- nated this plant as type of a new genus, Massowia, and published the combination M. cannaecfolia (Bot. Zeit. 10: 277-278). Apparently he was unaware that this species had been deseribed in the genus Spathiphyllum by Poeppig and End- licher (Nova Gen. Sp. Pl. 3: 85. 1845) as S. candicans.2 Massowia was placed in synonymy under Spathiphyllum by Schott in 1853, when he finally delimited his diagnosis of Spathiphyllum to exclude S. sagittaefolium? and expanded it to include related genera of other authors. Koch in 1856 (Bonplandia 4: 10-12) published a sharp rebuttal of Schott’s nomenclatural action. | Schott, in his Genera Aroidearum (1858), described Spathiphyllum and pre- sented a full-page plate of the reproductive structures of S. blandum. Two years later he presented his Prodromus Systematis Aroidearum, a monograph of the entire family with partial keys to the species, in which 22 species of Spathiphyl- lum were treated. Engler (Gard. Chron. II. 7: 140. 1877) erected the genus Amomophyllum, distinguished by the shape of the ovary and its uniovulate locules. In 1879 (in DC. Monogr. Plan. 2: 227) he altered the rank of Amomophyllum to that of a section of the genus Spathiphyllum. The last monograph of Spathiphyllum, prepared by Engler and Krause, ap- peared in Das Pflanzenreich (473°: 118-134. 1908). It was a good treatment, obviously representing new investigations undertaken since publication of the Monographie. Twenty-seven species and four varieties were treated. GENERIC RELATIONSHIPS The aroid genus Spathiphyllum is generally considered rather primitive.4 Its perfect flowers (with perianth) are borne on a spadix without a sterile ap- pendage, and the spadix is subtended by a more or less flat or cochleariform, persistent, rather foliaceous spathe. The geniculate apex of the petiole and the well-developed spathe are its only advanced characteristics, except in the few Species whose perianth is formed into a 4—6-sided, truncate cup. Engler and Krause (Pflanzenreich 478": 118-134. 1908) considered Spa- thiphyllum to be most closely related to Holochlamys because the perianth in the latter gefius is a 4-sided cup similar to that in S. commutatum. Holochlamys is similar in habit and foliage to Spathiphyllum, but is more advanced. Its ovary is unilocular with many ovules arising on long funiculi from a basal placenta, and generally the spathe is convolute about the spadix. Hutchinson (Fam. Fl. Pl. 2: 119. 1934) placed Holochlamys in the Lasieae near Podolasia because its spathe more or less embraces the spadix. On the con- trary, the unfurled and rather foliaceous spathe of Spathiphyllum led Hutehin- son to retain only this genus in the Spathiphylleac, which he considered to be more primitive than the Anthwrieae, and advanced only in relation to the Acoreae “8. candicans = 8. cannaefolium 3Schott ereeted the new genus Urospatha to receive S. sagittaefolium. 4The advanced conditions are: unisexual flowers (without perianth), and a spadix with a sterile appendage, enveloped in a well-differentiated spathe. 19601 SPATHIPHYLLUM 3 Fig. 1. Geographic distribution of Spathiphyllum. Numerals indicate the number of species in each area. Note the presence of a single species (S. laeve) on Cocos Island, and another (S. commutatum) in the Indonesian-Philippine area. and Orontieae. This treatment appears to be completely natural, and suggests that of Schott (Gen. Aroid. 1858), who placed Spathiphyllum between Anthurium and Orontium. Spathiphyllum is often confused with Stenospermatium and Rhodospatha. The latter two genera differ from Spathiphyllum in having flowers without perianth, a somewhat convolute spathe deciduous at anthesis, and a stem with elongate internodes. MORPHOLOGY The sections of Spathiphyllum are differentiated by (1) the discreteness or union of the perianth segments, (2) the form of the style and its length in relation to the perianth, and (3) the manner of attachment of the spathe upon the peduncle. Species differences are much less outstanding. The variation and the relative taxonomic merit of each structure are discussed below. Leaf-blade. The length/width ratio of the blade is a constant feature useful in differentiation of many species of Spathiphyllum, while the absolute size of the blade varies greatly. Shape of the blade is an important species characteristic, but the point at which the blade is widest varies somewhat. Texture of the blade is difficult to evaluate, especially in specimens glued firmly to an herbarium sheet, and is rarely considered in separating species. The base® of the leaf-blade may be acute or obtuse to subrotund, and is relatively constant within a species. It is noteworthy that the broad leaf-base 5The ‘‘base’’ of the leaf-blade here indicates the lower one-third or one-fourth of the blade where the leaf-margins converge toward the midrib; it should not be confused with the ultimate narrowing of the blade in some species into a small deltoid process. MEMOIRS OF THE NEW YORK BOTANICAL GARDEN Or 1960 | SPATHIPHYLLUM occurs mostly in species of sect. Spathiphyllum. The number of primary lateral veins and the angle at which they arise from the midrib seem to be constant features valuable in identification. Petiole. Though the petiole is similar in most species, the geniculum® may be exceptionally long or occasionally alate. Inflorescence. The length/width ratio and the shape of the spathe are essentially constant for a species. Texture of the spathe is rarely an important species characteristic. In most species, color of spathe varies with age of inflorescence. The spadix (as a unit) has little taxonomic value. In many species, it is borne on a free stalk, here called the stipe; this term always refers to the more or less terete, free apical portion of the peduncle between the point of attachment of the spathe on the peduncle and the base of the spadix. The length of the stipe is sometimes a good diagnostic characteristic. Flowers. The perianth furnishes adequate sectional differences only, except in two species in which the apex of the perianth is distinctively modified. This characteristic has not been previously recorded. In most other works on aroids, the perianth segments are designated as sepals or tepals. Since the 4-6(-7) perianth segments occur in a single series in Spathiphyllum, the terms ‘‘ perianth”’ and ‘‘perianth segments’’ are used throughout this paper. Specimens were examined in which the stamens appear to be of reduced size, but my study of these structures was inconclusive because of the small amount of material and the variability in the size of the stamens within a single flower (Fig. 2B). The pistil may be attenuate and exserted beyond the perianth, or inflated, annular, and/or truncate and scarcely exceeding the perianth. The proportional length of the exserted portion in respect to the length of the perianth may characterize a species, but more commonly a group of species, or a section. Ovary differences, relied upon heavily by Engler and others for separation of species of Spathiphyllum, are of limited taxonomic value in this genus. In some species the number of locules per ovary is reduced from the usual three to two, yet the bilocular situation may result by abortion only in some individual flowers of a spadix. In his key to the species, Engler emphasized the number of ovules per locule. My investigations have shown that this characteristic may be misleading. The number of ovules is sometimes inconstant from flower to flower and plant to plant. Moreover, the number of ovules in the locules of a single pistil is not equal; one locule tends to have one or occasionally two ovules more than the others. The largest number of ovules in any locule is significant, a smaller number simply indicating that a reduction has occurred. 6The geniculum is the thickened distal end of the petiole found in several genera of Araceae and in other families. Fic. 2. A-D. Reproductive structures typical of species of Spathiphyllum sect. Spathiphyl- lum. v S. humboldtii X S.gardneri e S. cannaefolium any SY 4 , g 2 Se we ae 9 a nae 2 iN ea oe le a) ee es v pA Agee, ( Se gee hy fs Mt lets \ : } mae je CAP el a Aig: ; $ . 1. oe } 1J Py vi . i } by 3 oe yee ee Cd. Bolivar Sy ol % OR g we { i 2 - 6 eats ~d2, =—.* ae > z Ts , = 4 AY i) 7 2 l- ee ee, A. pS eee, \ . | / " Ses ap fee : ¢ < wma. =) = es Maas i a eee a) Gr Hewr ay eee Sree f e see eS Ne S,grandi folium t ‘ A S.monachinol ae ee af Tees! \. & S.lanceaefolium A S. jejunum sas aN “it? = Vi S:kechi @ S.maguirei BS CORR yi a a a heneh e. 0 S. quindiuense V S.schomburgkii 300 soo mus | a sale m S.cuspidatum + S.neblinae a00- 500 600 KILOMETERS 3 rot 0 S.kalbreyeri @ S.sipapoanum Fic. 4. Geographic distribution of three species of Spathiphyllum widespread in northern South Ameriea. three species of sect. Spathiphyllum of northern South America and Panama. s Fig. 5. Geographic distribution of nine species of sect. Amomophyllum and they are typical. Most nearly identical to the type is Seibert 507, which has served to guide me in the amplification of this species. The Peruvian species S. lechlerianum is probably the closest ally of this species. From it.S. quindiuense is separated by its proportionally narrower spathe. S. quindiuense is considered primitive among the species of section Amomo- phyllum, and indeed may have arisen from stock similar to that which gave rise to S. friedrichsthaliui of sect. Spathiphyllum. Both species tend toward a multi- ovulate condition of the ovary, and both grow in the Panama-Colombia area where the greatest concentration of species of Spathiphyllum occurs. 1960 | SPATHIPHYLLUM Lee 5. S$. monachinoi Bunting, sp. nov. Figure 5. Foliorum lamina valde anguste oblonga ad lanceolato-ellipticam, 30-34 cm longa et 3.5-4.3 em lata, ad apicem acuminata, ad basim acuta, venis primarils 9- jugatis angulo 15-20° divergentibus; petiolus gracilis, 27-32 cm longus, ad dimidium conspicue vaginatus; geniculum ca. 1.7 em longum. Pedunculus peti- olum longitudine subaequans, ca. 30 cm longus; spatha anguste lanceolata, 11.8 em longa et 2.6 cm lata, apice attenuato, basi acuto et in pedunculo ca. 0.5 em decurrenti, intra viridi-alba, extus viridis (fide Wurdack et Monachino) ; spadix albus, ca. 4 cm longus, stipite ca. 2 cm longo; perianthii segmenta separata; pistilum obpyramidale, apice subtruncato, solum stigmate papilliformi peri- anthium superanti; ovarium 2-loculare, loculis 6—5-ovulatis, ovulis ad pulvinum subbasilarem in loculo quoque affixis, 12-11 in toto ovario; fructus ignotus. Type collection: J. J. Wurdack & J. V. Monachino 41056, ‘‘Spathe greenish- white above, green beneath, spadix white, in rocky moist ravine just below Raudal Maraca (ca. 110 river km from mouth), alt. 115 m,’’ Rio Parguaza, Edo. Bolivar, Venezuela; 31 Dee 1955 (holotype NY). Unicate. Known only from the type collection. The collectors told me that they encountered numer- ous sterile plants in the locality, but only one flowering plant was found and collected. It is with much pleasure that this species is named for Mr. J. V. Monachino, one of the collectors of the type specimen and Associate Custodian of the her- barium of the New York Botanical Garden. His keen taxonomic ability and willing assistance has greatly benefited the author during the course of this investigation. S. monachinot is the most robust Guayana species of sect. Wassowia. Its bilocular ovary contains twelve or eleven ovules per ovary, in contrast to the trilocular ovary of S. jejunum which contains a total of only seven to six ovules. S. monachinoi was collected in lower elevations along the Rio Parguaza, some 200 air miles north of the Rio Cunucunuma where S. jejunum was collected. Between these points lies the broad Rio Ventuari drainage, which Maguire and his asso- ciates have explored botanically without procuring a single specimen of Spa- thiphyllum. The greater number of ovules per locule in S. monachinoi easily sets it apart from S. sipapoanum from nearby Cerro Sipapo. It differs from the other species of Guayana by its outstandingly larger size and narrower leaves. There are similarities between this species and S. quindiuense. They differ in these characteristics: (1) the texture of the leaf-blade and spathe (in dried specimens) is chartaceous in S. monachinoi, pergameneous in S. quindiuense; and (2) the stipe of the spadix is proportionally shorter in the latter species. 6. S. gardneri Schott, Aroid. 1:2. pl. 3A. 1853. Figure 4. Leaf-blade oblanceolate to oblong-elliptic, (10—)14-34 em long, and 3-7.5.cm wide, the apex acute-acuminate, the base acute or subattenuate, with ca. 4-6 pairs of arcuate primary lateral veins arising at an angle of ca. 45°; petiole 15-35 em long, alate in the lower half or more; geniculum 0.8-2.4 em long. Peduncle exceeding the petiole, 21-45 cm long; spathe cucullate, lanceolate or elliptic, 7-10 em long and 2.2-3.4 em wide, the apex long-acuminate, the base acute to subtruncate, conspicuously oblique and decurrent on the pedunele to ca..1-em, white with a green central rib (fide Gardner) ; spadix 2.3-5.3 cm long, on a stipe ca. 0.5(-1.2) em long; perianth segments separate above, sometimes conglutinate below, at least in age; pistil obpyramidal,: scarcely exceeding the . 20 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [ vou. -10 perianth, apically conic-subtruncate, constricted between the annular style and the ovary, the stigma coronate; ovary 3-locular, the ovules superposed, varying from 6-3 in each of the 3 locules; totaling 15-11 ovules per ovary; fruiting spadix somewhat tuberculate; fruit obovoid, rostellate; seeds as many as the ovules, + oblong in profile and very angular, the inner faces smooth, the outer face emooth or somewhat verruculose along the angles. Type collection: G. Gardner 1867, ‘‘about 2 feet high, banks of the river near Crato; spathe white with a green genteel rib,’’ Ceara, Brazil’ Nov 1838 (lecto- type K, isotypes BM, G-BOIS, G-DC, G-DEL, 8, US). Distribution: River banks in the forests of Maranhao and Goias, Brazil. BRAZIL Cear4é: Gardner 1868 (P); Serra do Araripe, Procendencia do Granjeiro, Sep 1933, Luetzelburg 25842 (M). Goias: region of southern Serra Dourada at long. 48°50’W. lat. 13°45’ S, forest along road 13 km east of Formosa, Mai 1956, Dawson 15135 (LAM). Maran- hao: Don 2 (BR). This is the southernmost species of Spathiphyllum in eastern South America. For nearly a century S. gardneri was known from the type collection only. I am the first to assign Don 2 to this species. Though Lanjouw included G. Don among the West Indian collectors only, there can be little question that the specimen in the Martius Herbarium that bears his name was collected in Brazil and repre- sents S. gardnert. The original set of Don’s collections is at Edinburgh; un- fortunately, I have not seen that material. Two modern collections are identified as S. gardnert. Although Luetzelburg 25842 was collected in the type locality, the leaves are very small in comparison to those of other specimens of this species. Nearly identical to the type material is Dawson 15135, collected in the province of Goias. Schott originally described the sepals of S. gardneri as connate; Engler de- seribed the perianth as 6-lobed apically with the segments connate pelos which is essentially accurate. Such a criterion, however, is no good reason fein their placing this species in sect. Massowia with those species which have a cup-like perianth. S. gardneri appears to be closely allied to S. cuspidatum, and is in- cluded in sect. Amomophyllum. From the latter species, S. gardneri differs in its shorter spadix which is nearly sessile, and in the ovules which are superposed and affixed well up in the locules. 1-8: cuspidatum Schott, Oesterr. Bot. Wochenbl. 7: 158. 1857. Figure 5. Leaf- blade lanceolate to oblong-elliptic, 24-38(—51) em long and 5—10(-13) cm wide, the apex acuminate, the base acute, with ca. 9 pairs of primary lateral veins ene at an angle of 35-45°; petiole somewhat longer than the blade, (11—)25-45(-66) em long, conspicuously alate in the lower two-thirds; genicu- lum 1.2—2(-3) em long. Pedunele ca. 2—2.5 times as long as the blade, 35-70 (-100) em long; spathe lanceolate to elliptic, 10-20(-26) em long and 3-6 cm wide, the apex ‘dGurieaee to subeaudate, the base acute or narrowed into a short handle; spadix 4—-9(-13) em long, on a stipe 1—2.5(—3.8) em long; perianth of | separate segments; pistil obpyramidal, apically truncate, slightky constricted between the + annular style and the ovary, only the nipple-like stigma exceed- ing the perianth; ovary 3-locular, the ovules affixed near the base of the locules, varying from 4-2 in each of the 3 locules, totaling 12-6 ovules per ovary; fruit- ing spadix smooth; fruit obovoid, apically truncate; seeds as many as the ovules, oblique-oblong-ovate in profile, the surface quite smooth or shallowly furrowed and foveolate. : 1960] SPATHIPHYLLUM 21 Type collection: C. C. Parker s.n., ‘‘Demerara,’’ British Guiana (lectotype K). Apparently a unicate, collected in 1824. A Splitgerber collection from Suriname was also cited by Schott in the original publication of this species, but it has not been seen, or even located. Distribution: Swampy forested areas of British Guiana. BRITISH GUIANA: Mazaruni R., Appun 230 (K); Appun 848 (K); Smote Creek, Waini R., Jul 1906, Beckett § Kortright 8487 (K); Kaow Creek, Mazaruni Station, Jul 1939, B. G. For. Dept. 2976 (K) ; Potare, Mahdia R., Jan 1943, B. G. For. Dept. 3904 (K), 3909 (K) ; in swamp, Huradaiah, Moruka R., Pomeroon Dist., Aug 1921, de la Cruz 1026 (GH, NY, US); Pomeroon R., Pomeroon Dist., Jan 1923, de la Cruz 2940 (F, GH, MO, NY, US); Pomeroon R., Dee 1922, de la Cruz 3128 (F, GH, US); Waini R., Northwest Dist., 8°20’ N, 59°40’ W, Apr 1923, de la Cruz 3610 (F, GH, MO, NY, US); along creek, Kartabo Station, junction of Mazaruni R. & Cuyuni R., Jul 1924, Graham 310 (NY); swamp in forest by Kawow Creek, Penal Settlement, Dee 1919, Hitchcock 17235 (GH, NY, 8, US); Hooker sn. (NY); Pomeroon R., Mar 1884, Jenman 1985 (K); Demerara R., Jan 1896, Jenman 6868 (K); Barima R., Mar 1896, Jenman 6989 (K); Kamakusa, Nov 1922, Leng 49 (NY); Kamakusa, Dec 1922, Leng 353 (NY): wet places, Comaka, Demerara R., Mai 1923, Peraud 269 (F); in mora swamp, Moraballi Creek, near Bartica, Essequibo R., Aug 1929, Sandwith 153 (K). Engler and Krause (Pflanzenreich 4738: 128. 1908) cited Jenman 6989 as S. cuspidatum, but incorrectly stated its locality as ‘‘Roraima.’’ It is a mixed collection from along the Barima River; the New York sheet represents S. cannaefolium, the two Kew sheets S. cuspidatum. Eneler described the locules of the ovary of this species as biovulate; my observations are that there are 4-2 ovules per locule. S. cuspidatum appears to be limited to British Guiana. However, Jonker & Jonker (Fl. Suriname 1: 28, 29. 1953) included this species in their treatment on the basis-of a Splitgerber collection from Sarumassa, cited by Schott (Prodr. Syst. Aroid. 428. 1860). Although no subsequent investigator appears to have seen that specimen, it is possible that S. cuspidatwm occurs in Suriname. S. cuspidatum may represent a terminal element in phylogenetic develop- ment in this genus. It varies very little, and is common within narrow geo- graphical limits. It is most similar to S. kalbreyeri, from which it differs in its lanceolate or elliptic spathe which is somewhat broader than in the latter species. There is also a tendency toward a slightly greater number of ovules per ovary than is found in S. kalbreyeri. Of considerable interest is the possible relationship between S. cuspidatum and S. humboldti, which latter species exhibits such diverse ovary characteris- tics as to be placed in a separate section. It is a reasonable conjecture that these two species, as well as S. gardneri, may have arisen from a common primal stock. 8. S. kalbreyeri Bunting, sp. nov. Figure 5. Foliorum lamina e oblongo- ad lanceolato-ellipticam, 33-40 em longa et 9-10.5 em lata, ad apicem acuminato-cuspidata, ad basim acuta; venis primariis multis angulo 35—-45° divergentibus; petiolus 32-43 em longa, ad 14 vel plus vaginatus; geniculum ca. 2 em longum. Pedunculus 40-58 em longus; spatha oblonga, 13.5— 15 em longa et 2.5-3 em lata, apice attenuato-acuminato, basi acuto et leviter complexo; spadix 7—8.5 em longus, stipite ca. 1 em longo; perianthii segmenta Sseparata; pistillum obpyramidale, apice truncato, solum stigmate papilliformi perianthium superanti; ovarium 3-loculare, loculis 4~-2(—1)-ovulatis, ovulis ad pulvinum subbasilarem in locule quoque affixis, 9-5 in toto ovario; fructus ignotus. ; ‘ : 22 MEMOIRS ‘OF THE NEW YORK BOTANICAL GARDEN [vou. 10 Type collection: Kalbreyer 1413, ‘‘tufted habit, 214-3’; leaf erect, thin; spathe 5-6” 1., green, nerved when old inside white; spadix bent greenish or yellowish ; forest shady marshy spaces 3000’; Rastrojas,’’ Colombia, 15 Feb 1880 (holotype Kk). COLOMBIA: Sur de Santander: Amarilla Creek, vic. of Barranea Bermeja. Magdalen Valley between Sogamoso and Carare Rivers, Jul 1936, Haught 1894 (US). PANAMA: Darién: trial between Paya & Pucro, wet forested area, Jun 1959, Stern, Chambers, Dwyer & Ebinger 404 (MO). S. kalbreyeri is limited to the Magdalena Valley of Colombia and adjacent Darien, Panama. It is remarkably similar to S. cuspidatum, but is probably distinct. Each of these species has a very limited range, and is separated from the other by all Guayana and the Cordillera Oriental of the Andes. S. kalbreyeri is distinct in its oblong spathe which is proportionally somewhat narrower than in S. cuspidatum. There is a tendency toward a small reduction in the number of ovules per ovary in this species, and the blade of the leaf and the spathe are coarser in texture than in S. cuspidatum. 9. S. lechlerianum Schott, Prodr. Syst. Aroid. 425. 1860. Figure 6. Leaf-blade oblique, narrowly lanceolate-elliptic to oblong-elliptic, 16-25 em long and 2.5-4 em wide, the apex attenuate-acuminate, the base attenuate or acute, the 5 or 6 pairs of primary lateral veins arising at an angle of 25—-30° ; petiole 15-25 em long, prominently alate in the basal two-thirds or nearly to the geniculum in reduced floral leaves; geniculum 1.3-1.8 em long. Peduncle 27-42 em long; spathe cucullate, broadly lanceolate, 7-12 em long and 2.2—3 em wide, the apex acuminate to subeaudate, the base subacute and decurrent on the pedun- cle to 1.4 em (!), white within, medium green outside (fide Spruce); spadix 2—3.7 em long, on a stipe 0.5-1 em long; perianth segments separate or possibly conglutinate below in age; pistil obpyramidal, apically truncate, the elevated stigma scarcely exceeding the perianth; ovary 3-locular, the ovules superposed, varying from 5 to 3 in each of the 3 locules, totaling 14-10 ovules per ovary; fruit unknown. Type collection: W. Lechler 2338, ‘‘St. Gavan in umbros,’’ Peru, Jul 1854 (holotype K). According to Weberbauer (El Mundo Vegetal de los Andes Peruanos 14. 1945) this locality is San Gaban, in ‘‘el Oriente andino de la provincia de Carabaya,’’ which is northwest of Lake Titicaca and east or south- east of Cuzco. Distribution: Wooded slopes in southeastern and northeastern Peru, and the Magdalena Valley of Colombia (possibly). PERU: prope Tarapoto, Feb 1856, Spruce 4496 (BR, K). COLOMBIA: Sur de Santander: camp on Margarita Creek, vic, of Barranca Bermeja, Magdalena Valley, between Sogamoso R. & Colorado R., Sep 1934, Haught 1367 (US). CULT. (probably): Davis 152, ‘‘Hort. Veitch, Aug 1876, teste N. E. Brown,’’ and on same sheet an individual element labeled pene the Tee iE plants of Mr. Davis, coll. ‘Mess Veitch, ’?» Jul 1884 (K). This species extends almost to 14° S latitude. Specimens collected in widely disjunct localities have been identified as S. lechlerianum. The Colombian collec- tion (Haught 1367) must be referred to this species, despite the great distance between its collection locality and that of other specimens. However, it was not considered in the preparation of the description of S. lechlerranum. 1960 | SPATHIPHYLLUM 23 The original description of S. lechlerianum gives no details of the ovary, but Schott placed this species in a group which he characterized as having the sepals connate in a truncate cup! Engler retained this species in sect. Massowia, but emended the description of that group to account for the six-lobed nature of the apex of the perianth found in this species. N. E. Brown later annotated the type sheet ‘‘Perianth segments all free to the base.’’ I made careful examination of this structure in the type specimen and in Davis 152, and concluded that the perianth segments are free from one another. It appears that the segments may be conglutinate in age as in many other species. S. lechlerianum is closely allied to S. quindiuense, and is here included in sect. Amomophyllum. Though the locules of the ovary are characteristically 5—4-ovulate in both these species, the leaf-blade of S. lechlerianum is propor- tionally narrower and its lanceolate spathe is proportionally wider and of more delicate texture than that of S. quindiuense. From S. jejunum this species differs in the greater number of ovules per locule. 10. S. jejunum Bunting, sp. nov. Figure 5. Foliorum lamina obliqua, anguste oblongo-elliptica, 18-20 em longa et 2-2.5 em lata, ad apicem attenuata, ad basim acuta, venis primarlis 4-5-jugatis angulo 15-20° divergentibus; petiolus 10-14 em longus, ad 24 vaginatus; geniculum ca. 0.6 em longum. . Pedunculus 16 em longus; spatha angustissime elliptica, 5.8 «m longa et 1 em lata, apice attenuato, basi acuto, albo (fide Curran) ; spadix 1.6 em longus, stipite ca. 1 cm longo; perianthii segmenta separata; pistillum cylindrico- obpyramidale, apice obtuso-subtrunecato, solum stigmate elato perianthum super- anti; ovarium ut videtur 2-loculare, loculis 3—2-ovulatis, 7-6 in toto ovario; fructus ignotus. Type collection: H. M. Curran 226a, ‘‘found growing on rock along river banks, white flowers, Jacare, Rio Cunucunuma,’’ Terr. Amazonas, Venezuela, “4. 6. 1950’’ (holotype NY). Unicate, presumably. Known only from the type collection. The two plants that represent S. jejunum were found mixed in a collection of S. cannaefolium. This is the only collection representing sect. Amomophyllum from the region near Cerro Duida. It is likely that they were collected at a low elevation, though the altitude was not stated on the label. S. jejunum is related to S. monachinoi, but differs in its much smaller size (which suggested the spe- cific epithet), and the 2—3-ovulate locules of the ovary are very different from the multiovulate locules of S. monachinoi. The proportionally narrower leaf- blade and spathe and the shorter geniculum serve to differentiate this species from S. sipapoanum. From S. maguirei, S. jejunum can be separated by its narrower leaf-blade and its apparently persistent spathe. 11. S. maguirei Bunting, sp. nov. Figure 5. Foliorum lamina lanceolata, 13-15 em longa et 2.3-3 em: lata, ad apicem at- tenuato-acuminata, ad basim acuta, venis primariis ca. 4~jugatis angulo ca. 25° divergentibus ; petiolus 9.5-12 em longus, ad 34 vaginatus; geniculum 0.6—0.9 em longum. Pedunculus ad 23 em longus; spatha anguste lanceolata, ut videtur mature deciduo, 5.2-5.6 em longa et 1 cm lata, apice attenuato, basi late acuto, viridis (fide Maguire) ; spadix 1.5-3.3 em longus, stipite ca. 1 em longo; peri- anthii segmenta separata vel ad basi conglutinata vel connata; pistillum ob- pyramidale, apice subtruncato, solum stigmate elato perianthium superanti; ‘ 7 24 MEMOIRS: OF THE NEW YORK BOTANICAL GARDEN [vou. 10 ovarium 3-loculare, loculis 3—2-ovulatis, ovulis ad pulvinum subbasilarem in loculo quoque affixis, 7-6 in toto ovario; spadix fructiferus plus minusve levi; fructus angulari-sphaeroidens, latitudine longitudinem subaequanti, apice trun- eato; semina ovulos numero aequantia, obliquo-ovoidea, superficie foveolato. Type collection: B. Maguire & D. B. Fanshawe 32469, ‘‘terrestrial, spathe green; spadix green; on rocks in stream, Moruka Creek, locally common, 1200 m alt., Kamarang River, Wenamu Trail,’’ British Guiana, 9 Nov 1951 (holotype NY, isotype US). Known only from the type collection. Though S. maguire: occurs in eastern Guayana, it seems as closely related to the western Venezuelan species (S. neblinae and S. sipapoanum) as to S. schom- burgku from nearby Mt. Roraima. The lanceolate leaf-blade of S. magwirei is distinetly different from the narrowly elliptic blade of S. schomburgkw. S. maguirer finds its closest relative in S. neblinae, from which it is geo- graphically separated by some 500 air miles. The spathe is often deciduous in these species and their floral characteristics are similar. S. magwirei differs by its proportionally narrower leaf-blade which is widest below the middle, and in the shorter stipe of the spadix. No Spathiphyllum has been found in the intermediate area of great sandstone mountains of Bolivar to the west of Mt Roraima. 12. S. schomburgkii Schott, Oesterr. Bot. Wochenbl. 7: 158. 1857. Figure 7. S. candolleanum Schott var. schomburgku (Schott) Engl. in DC. Monogr. Phan. 2: 225. 1879. Leaf-blade oblique, narrowly elliptic, to 18 em long and 2.5 em wide, the apex acute-acuminate, the base attenuate, primary lateral veins ca. 4 pairs arising at an angle of 15-25°; petiole to 12.5 em long, prominently alate nearly to the geniculum ; geniculum ea. 0.8 em long. Peduncle 26 em long; spathe lanceolate- elliptic, 6.2 em long and ea. 1.6 em wide, the apex acute-acuminate, the base acute; spadix ca. 2.5 em long, on a stipe ca. 0.7 em long; perianth of separate segments; pistil obpyramidal, scarcely exceeding the perianth, the apex obtuse- subtruneate, the stigma coronate; ovary 2-locular(?), the ovules affixed on a sub-basal pad in each locule, (apparently) 4-3 in each of the locules, totaling 7-6 ovules per ovary; fruit unknown. Type collection: R. H. Schomburgk s.n., ‘‘1st Coll. M. Roraima,’’ Venezuela (holotype K). Unicate(?). Probably found at lower elevations on slope of Mt. Roraima. Known only from the type collection. S. schomburgkwui stood for a long time as the only known species of this genus from a sandstone mountain of Guayana. Subsequent exploration of this region has yielded several collections of Spathiphyllum. All represent plants of small size. The pistil is nearly identical in all, viz., cylindrical-obpyramidal and scarce- ly exceeding the perianth, the apex obtuse-subtruncate, the stigma nipple-like or coronate. The pistil of S. schomburgkwu was described initially as attenuate with a conical apex exceeding the sepals. On this basis (apparently), Engler made it a variety of S. candolleanum. Considerable attention to the nature of the pistil of this species has led me to conclude that it is unquestionably similar to that in other collections from sandstone areas of Guayana. It is difficult to determine the number of taxa represented among the Gua- yana collections. In view of the small number of collections, I was reluctant to create new names that might later fall in the light of further collections. Never- theless, it is felt that no collections are conspecific, and six species are described as new to science. : 1960! SPATHIPHYLLUM Pay S. schomburgkti seems to be closely related to 8S. sipapoanum from which it differs by its narrowly elliptic leaf-blade, its perianth of five or six segments," and the greater number of ovules per ovary. It is separated by its narrower leaves and the shorter stipe of the spadix from SN. neblinae. Inflorescence charac- teristics serve to set apart S. schomburgkui from the Andean species. 13. S. neblinae Bunting, sp. nov. Figure 5. Foliorum lamina elliptica vel rare lanceolato-elliptica, (10.5—) 15-19 em longa et (2.3-)2.9-3.9 em lata, ad apicem acuminata, ad basim acuta, venis primariis 4-5-jugatis angulo 20—-30° divergentibus; petiolus (4-)7.5-12 em longus, ad dimidium vel plus vaginatus; geniculum 0.5-1.3 em longum et supra sulcatum. Peduneulus (6—)18-25 em longus; spatha recta, e oblonga ad lanceolatam, (4.4-)5.7-6.8 em longa et (0.9-)1.3-2.4 em lata, apice attenuato-acuminato, basi attenuato et aliquando breviter ansato, albo florescenti, in aetate viridescenti et ut videtur deciduo ; spadix viridis, (1—)1.8—3.8 em longus, stipite (1.2—)1.7-2.4 em longo; perianthii segmenta separata; pistillum obpyramidale, apice truneato, leviter constrictum inter ovarium annulare et stylum, solum stigmate elato perianthium superanti; ovarium ut videtur 2-loculare, loculis 5—2-ovulatis, ovulis ad pulvinum subbasilarem in loculo quoque affixis, 9-5 in toto ovario; spadix fructiferus levis; fructus cylindrico-obpyramidalis; semina ovulos numero aequantia, valde angularia et irregulariter reniformia vel obliquo-ovoidea, super- ficie conspicue foveolato. Type collection: B. Maguire, J. J: Wurdack & G. 8S. Bunting 36797, ‘‘spathe white in fir., green in frt., spadix green; geniculum grooved above; rainforest: streamside at Camp 2; locally frequent; elev. 180 m; lowland & slope forests. Cerro de la Neblina, Rio Yatua’’; Terr. Amazonas, Venezuela; 14 Dee 1953 (holotype NY, isotype US). VENEZUELA: Maguire et al. 36799, same data as the type collection (NY). S. neblinae is allied to the other species of Guayana (discussed under SV. schomburgkit). Its closest relative is S. maguirei. Both share the peculiarity of a deciduous spathe, but from all other species of the sandstone area S. neblinae differs by its proportionately wider leaf-blade and the longer stipe of its spadix. It resembles S. minor of Amazonian Peru, but the latter differs in the shorter stipe of its spadix and the uniovulate condition of the locules of its ovary. 14. S. floribundum (Lind. et André) N. E. Br. Gard. Chron. IT. 10: 783. 1878. Figure 7. Anthurium floribundum Lind. et André, Ill. Hortic. 21: 24. pl. 159, 1874. Amomophyllum floribundum (Lind. et André) Engler, Gard. Chron. TI. 7: 140. 1877. Spathiphyllum floribundum var. cuneatum Engl. Bot. Jahrb. 6: 281. 1885. Leaf-blade conspicuously oblique, elliptic to oblong or oblanceolate, widest at or just above the middle, 13—20(—26) em long and 5.5—-9(-10.5) em wide, the apex cuspidate, the base obtuse or acute, marginally undulate, dark green above with a velvety lustre, the primary lateral veins ca. 9 pairs arising at an angle of ea. 65-75° ; petiole (6.5—)10—22(-28) em long, often alate nearly to the geni- eculum, vaginate below or the wings expanded and free; geniculum 0.5-1.3 em long. Peduncle (16—)20-37 em long; spathe white, reflexed, lanceolate to oblong- elliptic, 4+-8(—9.8) em long and 1.2—3 em wide, the apex attenuate to euspidate, the base acute to subtruneate and clasping the peduncle; spadix 2.5-5.5 em long, on a stipe’ 0.3—0.8 em long; perianth green, the segments separate or sometimes 11INot considered a good diagnostic feature. 26 MEMOIRS’ OF THE NEW YORK BOTANICAL GARDEN [vou. 10 eonglutinate in age; pistil white, obpyramidal, apically truncate, the stigma elevated and shortly exceeding the perianth; ovary 3-locular, the ovules affixed at or near the base of the locules, varying from (3—)2 to 1 in each of the 3 locules, totaling (7—)6-4 ovules per ovary; fruiting spadix smooth; fruit spheroid, the apex truneate; seeds 1 or 2 per locule, the surface vertically furrowed and foveo- late between the verrucose ridges. Type: pl. 159, L’Ilustration Horticole 21: 24. 1874. This color plate ade quately defines S. floribundum, and agrees with the original description which it accompanies. This species was described from a plant growing in cultivation, and no type material appears to have been prepared. A fine representative specimen of S. floribundum is Lehmann 7758 (F, GH, US), which is more or less identical to the material cultivated today under this name. Distribution: Banks of streams in the central Magdalena Valley of Colombia. COLOMBIA: Boyaea: high forested fronts, El Humbo, 130 mi n of Bogoté, Mar 1933, Lawrence 696 (MO). Cundinamarea: Paime, Bro. Ariste-Joseph A924 (US). Sur de Santander: camp on Taza Creek, vicinity of Barranca Bermeja, Magdalena Valley between Sogamoso R. & Colorado R., Feb 1934, Haught 1271 (US); camp on Margarita Creek, vicinity of Barranea Bermeja, Magdaiets Valley between Bngninole R. & Colorado R., Cet 1934, Haught 1386 (F, US); brookside, Dosquebrados, Jan 1880, Kalbreyer 1354 (K). Antioquia: Rostrojas, Feb 1880, Kalbreyer 1414 (K); Sabaleta, Lehmann 7757 (F); Rio Nuz, 600-1000 m, Lehmann 7758 (F, GH, US); ‘‘alt. 1200,’’ Triana 696 (BM). CULT.: Hort. Williams, Aug 1876, and Hort. Kew, Mai 1876, V. E. Brown s.n. (K); Hort. Kew ex Hort. Williams, Apr 1878, and Hort. Veitch, Apr 1879, NV. E. Brown s.n. (K); Engler 82 (B, BM, C, G-BOIS, G-DEL, GH, L, M, P, US); Hort. Cantonspaark no. 3393, Dee 1953 (U); ete. Perhaps the most handsome member of the genus, this species has been a eae conservatory subject for nearly a century. Though not described until 1874, S. floribundum was first collected by Gustay Wallis before 1864. The ca description gave no data on its arrival in Europe, but stated only that it was prepared from a plant growing in the garden of M. Linden for the previous two years, and that its publication had been delayed by lack of sufficient flowers to determine in which genus it should be placed (they erroneously selected Anthurium!). In 1875 (Hamb. Gart. Blum. 31: 63), Wallis listed A. floribundum among his collections near San Carlos (Aguas Claras), New Granada, mentioning that many of these same species were collected previously. It is likely that SN. floribundum was first collected in that locality. S. flor iundum appears to be endemic to the Magdalena Valley of Gotanae Various authors have referred to this species collections from Panama, but I cannot corroborate their identifications. Vegetatively Bailey 209 resembles this species, but a cursory ae of the spadix clearly demonstrates the conic pistils; it is referred to S. phryniifolium. Standley (Ann. Missouri Bot. Gard. 31: 34. 1944) cited this eaflcctan as S. floribundum, then illustrated the species with a drawing of Seibert 507, which I identify as S. quindiuense. There seems little cause to maintain var. cuneatum of this species. It was based upon Lehmann 2592, collected in Tolima, Colombia. That specimen was deposited at Berlin, and was probably burned. pe S. floribundum is outstanding for its dark green leaves with a velvety lustre. As a dried specimen, however, it differs in its broad leaves. It is most closely re-— lated to S. fulvovirens and S. juninense, both of which are species of much greater size than this species. Some cultivated material of S. floribundum has the spathe cuspidate at the Io 4 1960} | ; - SPATHIPHYLLUM apex, in contrast to the long-attenuate apex of the spathe in all collections from the wild. This difference of the cultivated form may have resulted from selection by horticulturists. 15. S. fulvovirens Schott, Oesterr. Bot. Zeitschr. 8: 179. 1858. Figure 7. Leaf-blade oblique, typically ovate to ovate-elliptic, (25—) 35-44 cm long and (10—) 17-20 em wide, the apex terminated by a cusp 1.5—3.5 cm long, the base ob- tuse to subrotund, the many primary lateral veins arising at an angle of 60—70° and very prominent beneath; petiole to 74 cm long, alate in the basal two-thirds ; geniculum (1—)1.5-3 em long. Peduncle to 49 em or more long; spathe + oblong and clasping at the base; spadix 6.8-11.5 em long, smooth, on a short stipe 0.3—0.8 em long; perianth of separate segments; pistil obpyramidal, apically truncate, the nipple-like stigma scarcely exceeding the perianth; ovary 3-locular, the ovules affixed near the base of the locules, 2 or 1 in each of the 3 locules, totaling 6—5 ovules per ovary; fruit unknown. Type collection: H. von Wedel 2198, ‘‘shrub 3 ft., inflorescence gray, Fish Creek, vicinity of Chiriqui Lagoon,’’ Prov. Bocas del Toro, Panama, 9 Apr 1941 (neotype GH, duplicates F, MO). The designation of a neotype is appropriate since neither the holotype (Wendland 939, ‘‘Pedregal, C. Rica, 1857’’) nor any dupheate of it has been located. A photograph of the holotype (Field Mus. no. 29838) clearly indicates that the type was deposited at Vienna, and therefore we must assume that it was burned along with the other Araceae. Distribution: {Pedregal(?), Costa Rica], along Chiriqui Lagoon in northern Panama, in the central Magdalena Valley and Pacific coast of Colombia (Buena. ventura). COLOMBIA: Rio Magdalena, Weir 55 (BM). Santander: Amarilla Creek, vicinity of Barranca Bermaja, Magdalena Valley, between Sogamoso R. & Carare R., Jul 1936, Haught 1895 (F, US)~ Valle del Cauca: Rio Dagua, Buenaventura, Lehmann 5361la (F, K). CULT.: ‘‘coll. Fl. Claes, Colombia, Juin 1927,’’ V. Lambert s. n. (BR). A good photograph of the holotype in conjunction with the original descrip- tion of S. fulvovirens serves to define this species very well. It is further elari- fied by an original Schott drawing (W). The exact collection locality of Wendland 939 is uncertain. Standley (Field Mus. Publ. Bot. 18: 48. 1937) noted that Wendland entered Costa Rica ‘‘by the Sarapiqui Valley, and explored especially the mountain chain from Barba to Turrialba,’’ which would limit his localities to the provinces of Heredia and Cartago, on the Atlantic slope of the Cordillera. The Colombian collections referred to this species have a somewhat narrower leaf-blade than is typical, but they are essentially similar in other features. S. fulvovirens finds its nearest relative in S. juninense, a Peruvian species with a wider spathe. These two might well be considered as infraspecific categories of a single species, but are maintained here as separate species because of the great distance between their areas of collection and the natural barrier to continuity of distribution created by the Cordillera Oriental of the Andes in Colombia. 16. S. juninense Krause, Notizbl. Bot. Gard. Berl. 11: 615. 1932. Figure 6. Leaf-blade oblique, ovate-elliptic, 28-32 em long and 12-15.5 em wide, the apex cuspidate (probably; lost from type specimen), the base rotund-obtuse, with many primary lateral veins arising at an angle of ca. 55-60° ; petiole 40-57 em long, alate throughout much of its length; geniculum 1.6—2 em long. Peduncle to 70 em long; spathe oblong, to 19 em long and 5.7 em wide, the apex long-cuspi- date, the base apparently obtuse; spadix to 12.3 em long, on a stipe ca. 1.5 em long; perianth of separate segments; pistil obpyramidal, the apex subtruncate, 28 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [vou. 10 © @S. friedrichsthalii s S. wendlandi: ‘< x» S. atrovirens a S. phryniifolium Vv S. floribundum © S. fulvovirens C Oe 6 Purse x S. lechlerianum V S. juninense @ S.tenerum O S.minor wm S.gracilis A S.patulinervum 100 200 300 400 MILES 100 200 300 400 soo 600 MILES oy, er 4 = 1 + —_ : - 2