A New Subsection of Anthurium Section Calomystrium (Araceae) and Five New Species from Colombia and Ecuador

Home , Anthurium

T. B. CROAT,]. WHITEHILL, E. D. YATES, 2007 23 A New Subsection of Anthurium Section Calomystrium (Araceae) and Five New Species from Colombia and Ecuador

Thomas B. Croat Missouri Botanical Garden P. O. Box 299, St. Louis, MO 63166-0299 [email protected] Jane Whitehill 309 E. 108th St., Apt. 5G New York, NY 10029 [email protected] Emily D. Yates Millennium Seed Bank Project, Co·coordinator Institute for Plant Biology and Conservation Chicago Botanic Garden 1000 Lake Cook Road Glencoe, IL 60022 [email protected]

ABSTRACT Calomystrium, owing to their rupicolous habitat. Subsection Rupicola, a new subsection of Anthurium section Calomystrium is proposed, encompassing nine rupicolous KEYWORDS species. Descriptions and a key are pro Araceae, Anthurium, section Calomy vided for these species including: A. strium, Colombia, Ecuador, Panama, new antrophyoides Killip, from Ecuador and species, taxonomy, new subsection, sub Colombia, A. chocoense Croat and A. section Rupicola. antioquiense Engler from Colombia, as well as A. amnicola Dressler, and A. INTRODUCTION sytsmae Croat from Panama. Five species, A. palacioanum Croat and A. weiffii In his revision of Anthurium for Panama Croat from Ecuador, and A. chocoense (Croat, 1986), the senior author recognized Croat, A. callejasii Croat, and A. vander an unusual group of rare species occurring knaapii Croat, from Colombia, are de only on rocks in rapidly moving streams. scribed as new. Members of this group These species were first believed to belong were previously thought to belong in to section Porphyrochitonium despite the section Porphyrochitonium because of lack of foliar glandular punctations typical their generally lanceolate leaf blades, of Porphyrochitonium. This group of spe and stems with short internodes. Howev cies shared with section Porphyrochito er, lack of distinct glandular punctations nium their small stature, short, slender on blade surfaces and the ability to internodes, and lanceolate to oblong-ellip interbreed readily with other members of tic blades. It was therefore assumed that section Calomystrium indicate that this these species were members of section new group belongs in section Calomy Porphyrochitonium that had lost their strium, and has evolved without conspic glandular punctations. Of this group dis uous posterior lobes typical of section cussed in the Panama revision (Croat, 24 AROIDEANA, Vol. 30

1986), only Anthurium rupicola Croat has group of rupicolous species, all believed to proven upon further investigation to be an be rare, arose from the more typical actual member of A. section Porphyrochi members of the section by progressively tonium. Anthurium rupicola is the only adapting to a rheophytic life form along species in section Porphyrochitonium thus and eventually into the bed of streams, far, that takes on a rupicolous habit. where they occur on rocks, much like However, because of their streamlined Spathiphyllum quindiuense Engler. This shape, more species in section Porphyr group is sufficiently distinct from other ochitonium could be expected to be more typical members of section Calomy rupicolous owing to the fact that they could strium with prominently cordate leaf grow in swiftly moving water without blades to warrant subsectional recognition. being dislodged. Breeding studies by H. The name Rupicoia meaning "thriving Kamemoto (pers. comm.) and others have among or inhabiting rocks", is proposed shown that some typical members of the for this subsection. The terminology and small rupicolous group discussed here, descriptive terms in the present paper e.g., Anthurium antioquiense and A. am follow (Croat & Bunting, 1979). The new nicola, readily interbreed with typical subsection Rupicola is characterized as members of section Calomystrium such as follows: A. andreanum Andre. This offers strong evidence that species comprising this small RuPicoia Croat, subsect. nov. TYPE: An rupicolous group are actually members of thurium antioquiense Eng!. Das Pflan section Calomystrium that have lost the zenr. IV. 23B. 174. 1905. cordate blades with prominent posterior lobes that are typical of this section. Differt a sect. typica per lamina plus Anyone who has tried to carry a potted minusve lanceolata, habitu rheophyta. plant of typical members of sect. Calomy A key to A. amnicola, A. antio strium (with their prominent posterior quiense, and A. antrophyoides was lobes that regularly hook themselves on provided by Dressler (980) and a key to other plants) down the narrow aisles of to the Panamanian species, A. amni a greenhouse will realize that such plants cola and A. sytsmae, was compiled by would not survive long on a rock in Croat (986). Below, these previous a rapidly moving stream. The tough poste keys are expanded to include all nine rior lobes act much like anchors, catching species in subsection Rupicola empha on anything with which they come in sizing the main differences among the contact. It is hypothesized that the small species.

KEY TO SPECmS OF SECTION CALOMYSTRIUM SUBSECTION RUPICOLA 1. Leaves with single midrib, either lacking a pair of basal veins or with a single pair of basal veins (one on either side of the midrib) promptly merging with the margin. 2. Spathe flamingo-orange; spadix orange...... A. vanderknaapii Croat. 2. Spathe lanceolate or elliptic, usually green, sometimes white or lavender-purple; spadix purplish, whitish or greenish. 3. Leaf blades more than 30 cm long, broadest well above the middle; collective veins arising from just above the base; tertiary veins not prominulous; Colombia, Antioquia, ca. 200 m ...... A. callejasii Croat 3. Leaf blades less than 22 cm long, broadest at about the middle;collective veins arising from one of the primary lateral veins in the middle or lower third of blade; tertiary veins prominulous; Central Panama and NE Ecuador, 200- 550 m. 4. Spadix green; berries early emergent, tapered and pointed at the apex, initially obovoid and green, maturing red-orange; cataphylls persisting as T. B. CROAT,]. WHITEHILL, E. D. YATES, 2007 25

thin, yellow-brown fibers; Central Panama, 200-400 m ... A. sytsmae Croat 4. Spadix yellow; berries emerging mature, globose and purple; fibers of weathered cataphylls very persistent; NE Ecuador, 500-550 m ...... A. weif.fi.i Croat 1. Leaves with 3 to 9 veins (including the midrib) at the base of the leaf blade and with one or more pairs of basal veins extending all the way to the apex. 5. Leaves rhombic or ovate-rhombic, broadest well below the middle; basal veins 7-11 veins at base of leaf blade (3-5 basal veins per side) or with basal veins lacking; spadix white. 6. Leaf blades rhombic" drying subcoriaceous, dark brown on upper surface; spathe white; Colombia, slopes of western Andes in Valle Department, 0- 250 m...... A. antrophyoides Killip 6. Leaf blades ovate-rhombic, drying moderately thin, greenish to grayish yellow-brown on upper surface (if the weak submarginal veins are included); spathe green; Ecuador, slopes of eastern Andes in Napo Department, 1800 m ...... A. palacioanum Croat 5. Leaves lanceolate or elliptic, with 3 veins at base ofleafblade (midrib and a single pair of basal veins); spadix white or lavender. 7. Primary lateral veins numerous, sometimes not distinguishable from interprimary veins; cataphylls weathering into persistent fibers; spathe green; spadix white at anthesis...... A. chocoense Croat 7. Primary lateral veins 3-5 pairs per side; cataphylls deciduous with only their papery bases remaining or persistent and intact, never fibrous; spadix dark violet-purple, yellowish or pink at anthesis. 8. Spathe ovate, lavender to purplish violet; spadix dark violet- purple; flower scent spicy, but not smelling of mint; fruits almost purple; stems 4- 6 mm diam.; cataphylls soon deciduous, only their papery bases remaining; Central Panama, Cocle Province ...... A. amnicola Dressler 8. Spathe lanceolate, usually white, rarely lavender; spadix lavender; flower scent minty; fruits green or white; stems 12-15 mm diam.; cataphylls 5- 11 em long, persisting intact, drying reddish brown...... A. antioquiense Engler

Anthurium amnicola Dressler, Aroi spicuous; blades subcoriaceous, semi deana 3: 55. 1980. Anthurium lilaci glossy, elliptic to elliptic-lanceolate, gradu num Dressler non Bunting, Selbyana ally acuminate at apex, narrowly acute to 2: 300. 10 Jan. 1979 (Dec. 1978), non rarely obtuse at base, 7-14.5 X 1.2-3.7 em, Bunting, Dec. 1975. Type: PANAMA. usually broadest near the middle; upper Col6n: along tributary between Caiio surface sparsely and inconspicuously glan Rey and San Lucas, S of Cocle del dular-punctate; lower surface epunctate; Norte, 19 Aug. 1972, on boulders, 1.5- midrib acutely raised above and below, 2 m above river, R. L. Dressler 4210 sunken toward apex above; basal veins 1 to (holotype, US; isotypes, MO, PMA, 2 pairs, weakly sunken; primary lateral SEL). Figures lA, lB. veins 3 to 4 per side, more or less obscure; Rupicolous; stems to 30 em long, 4- collective vein arising from the first basal 6 mm diam.; roots few, less than 2 mm vein, 3-4 mm from the margin. INFLORES diam.; cataphylls very thin, 4-5 cm long, CENCE erect; peduncle terete, 6-26 cm drying reddish brown, perSisting intact. long; spathe subcoriaceous, pale lavender LEAYES erect-spreading; petioles 6- (B & K purple 6/5), ovate, 2.6-4.2 X 1.5- 11 cm long, subterete, shallowly sulcate 2.2 cm, abruptly acuminate to cuspidate at adaxially; geniculum 1-2 cm long, incon- apex, rounded to attenuate at base, inserted 26 AHO IDEANA, Vol. 30

Fig. 1. A-B. Anthu riu111 a111nico/a Dressler. (SEL # 76-0053-002). A. Habit. B. Inflorescence, cl ose-up. C-D. Anthur iu m antioquiel1se Eng \. (H amme/ 15431A). C. Habit. D. Inflo rescence, close-up.

at 45° angle o n peduncle; sti pe 3-7 mm st

Fig. 2. A-B. A11fh u rium antrophyoides Killip. (Croat 56789). A. H abit. B. Inflo rescence, close-up. C-D . Al1lh uriu l17 callejasii Croa t. (Croal 77270) . C. Habit. D. Inflo rescence, close-up.

times waxy, elliptic, 1!-9.3 X 2.3-3 cm , AJ7lburiul17 anl rophyoides occurs in Co acute rounded at apex and abruptly lo mb ia (Valle and Ca uca) and Ecuado r ca udate-acuminate (acumen 1 cm lo ng); (Napo) at 100-400 Ill. stipe 1 cm long; spadix 1-3 X 0.5 cm , According to Engler's revisio n o f A n erect, w hite to crea m o r pale green; tburium in Das Pjla nzenreich , this sp ecies perianth segments equal , €I I out 0.8 mm is IllOS t similar to A. weberbauer i Eng!. in X 1 111111 . venation and general shape o f the leaves. 30 AROIDEANA, Vol. 30

The leaves of A. antrophyoides, however, lata; nervi primarii laterales obscuri; spatha are acute at base, not obtuse; the spathe is viridis; spadix atro-violascens-purpureus. proportionately much broader, and the Internodes 1-1.5 cm diam., shorter peduncles are shorter than the leaves, than broad; cataphylls persisting reddish while in A. weberbauri they exceed the brown and intact; petioles 17.5-26 cm X leaves, and the flowers are smaller. Com 2-2.5 (4) mm, drying slightly ribbed, light parison of the type specimen of A. antro green to medium brown; geniculum terete; phyoides with type material of A. weber blades 30-33 (-44) cm long, 3.5-4 (-5) cm baueri at Berlin (B) has been made by wide at widest point, widest just above the Killip. The leaves of A. antrophyoides also middle, linear oblanceolate, subcoria bear a very close resemblance to the fronds ceous, weakly glossy and slightly bicolor of the tropical African fern Antrophyum ous, drying grayish olive-green above, mannianum Hook. paler silvery green below; midrib convex paler and dark-green speckled above, Additional specimens seen-COLOM convex and moderately paler below; pri BIA. Valle: Cali-Buenaventura, km 104, mary lateral veins and collective veins Rio Tatabro, Idrobo & Kyburz 1769 (US); equally obscure, all arising at ca. 50 0 angle. Bajo Anchicaya, W A. Kyburz 2 (NY, UC, INFLORESCENCE erect-spreading; pedun US); vic. Santa Rosa, RIO Caballete, Killip cle 26-33 cm long, 1-1.5 mm diam.; 11555 (GH, US); Mun. Buenaventura, spathe green, spreading and recurled un Correg. Bendiciones, rd. to Aguaclara, der, drying medium brown to slightly Calima medio, Devia & Prado 2384 (MO); reddish brown, 4-6 mm wide, lanceolate; Correg. Cordoba, San Cipriano, Res. Nat. spadix green pre-anthesis, soon turning del RIo Escalerete, Devia et al. 3943 (MO); dark Violet-purple, matte, 3-7 cm long, RIO Agua Claro, ca. 40 km from Pacific drying medium brown to reddish brown, Ocean, Winters 9886 (US); ECUADOR. drying ca. 3 mm diam. Napo: RIO Granadilla, Codo Alto,Jaramillo Anthurium callejasii is known only from et al. 12681 (COL, HUA, QCA). the type locality. It is distinguished by linear-oblanceolate, green-drying blades Cultivated collections-COLOMBIA. with with a single pair of collective veins arising out locality, Croat 56789 (COL, JAUM, MO); somewhat above the base, and by the Cauca: Rio Timbiqui, Lehmann 8634 (F, green spathe and spadix. It can be com US). Valle: Buenaventura area, Selby 81- pared with A. systmae and A. we1jJii, both 2578, Plowman 14109 (SEL); Selby 81- of which differ in having shorter blades 2578, 25 Jan. 1986, Christenson 1150 (SEL). (less than 22 cm long), being broadest at ECUADOR. Without locality, cult. by Tom about the middle, and by having the Fennel, Florida, 1980, Fennel A (MO). collective veins arise from one of the primary lateral veins from the middle of Anthurium callejasii Croat, sp. nov. the blade to the lower third of the blade as Type: COLOMBIA. Antioquia: Mun. well as with prominently reticulate tertiary Taraza, Finca Callejas, ca. 200 m, veins. In contrast, A. callejasii has blades originally collected by Bruce McAlpin, broadest well above the middle with the received from New York Botanical collective vein arising just above the base of Garden 1383178c, vouchered 5 Oct. the blade, and obscure tertiary veins. 1994 as T. B. Croat 77270 (holotype, MO-04660293, isotypes, K, HUA, US). Anthurium chocoense Croat, sp. nov. Figures 2C, 2D. Type: COLOMBIA. Choc6: vic. of "La Equis" mine, short road N of Bollvar Internodia 1-5 cm diam.; cataphylla per Quibd6 Highway, around km 182-183, sistentia intacta; petiolus 17.5-26 cm longus, 300 m, 31 Oct. 1983, A.Juncosa 1175 2-2.5 (-4) mm lata; lamina linearis-oblan (holotype, MO-3158418, isotypes, B, ceolata, 30-33 (44) cm longa, 3.5-4 (-5) cm COL, JAUM, K, US). Figure 3A. T. B. CROAT, J. WHITEHILL, E. D. YATES, 2007 31

. , ..... "... - ,.. ,~,; .. ,,, ~- ,... ~,

...... <~H e, ~"~,,, .• r~ a""" .. ", .. . •.•• ,••.. "' .. "" I~"·U .. · ~."k. ' ...... ~~~l~ tmh::~-:'"~ : :::~ ~ ., .. ,,~

Fig. 3. A. Anthuriw17. chocoel1se Croat. (funcosa 1175). Herbarium type specimen. B. Anthu.rium palacioanum Croat. (Palacios 5580) . Herbarium type specimen . C. Anthu rium sylsmae Croat. (Davic!se & H amilton 23578). Herbarium type specimen. D. Anthurium vanderknaapii Croat. (Croat 903 41). Habit.

Herba rupicola; inte rnodia brevia, 8- primarii laterales numerosi, debil es; ped 15 mm diam.; cataphylla 5.5-6.5 cm unculus 30- 53.5 cm longus; spath a angu longa, persistentia in fibras solut a; pe ti o ste lanceolata , viridis, 3.5--4 cm longa, lus subteres, levite r sulcatus, 8-23 cm 5-8 mm lata; spadix 3.7- 6. 0 cm longus, longus; lamina folio rum oblo ngo-elliptica , 3- 6 mm cliam. , luteus, sessilis; baccae 21- 28 cm lo nga, 1.5-2. cm lata; nervi albae. 32 AROIDEANA, Vol. 30

Rupicolous herb; stems short; inter broadly rounded, outer margin obtusely nodes short, 8-15 mm diam.; roots numer 2-3-sided. INFRUCTESCENCE not seen, ous; leaf scars hidden by cataphylls; cata but berries white (fide Idrobo 1901). phylls 5.5-6.5 cm long, briefly intact, then Antburium cbocoense is endemic to weathering into fine, pale fibers; petioles Colombia, known only from the Depart subterete, weakly sulcate adaxially, 8- ment of Choco. All collections have been 23 cm long, 0.5-0.7 the length of the found either on rocks in streams or rooted blades; geniculum 8-15 mm long, drying at the margins of streams. Flowering plants darker than surface; blades subcoriaceous, have been collected in February, April and narrowly oblong-elliptic, gradually acumi July. nate at apex, narrowly acute at base, 21-28 The species is easily confused with X 1.5-2.7 cm, broadest usually at the Anthurium rupicola Croat, a species from middle; upper surface dark green and Panama in section Porphyrochitonium iridescent, lower surface yellow-green, having remarkably similar features includ both surfaces drying matte to weakly ing the size and shape of blades, spadix and glossy, epuncate on the upper surface, spathe color, as well as similar persistent inconspicuously and densely pale-punctate pale cataphylls. However, A. chocoense on the lower surface (punctations irregu lacks the typical glandular-punctate lower larly shaped, raised, but not apparently blade surfaces characteristic of section glandular); midrib convex and concolor Porpbyrochitonium. Instead, A. chocoense ous above, drying prominently raised, has irregularly shaped, pale, somewhat narrowly raised and paler below, drying raised, non-glandular punctations. Anthur raised and irregularly ridged; primary ium chocoense also differs from A. rupicola lateral veins many, not at all distinguish in having sessile, rather than the usually able from the interprimary veins upon stipitate spadices of A. rupicola, and drying, departing midrib at 20-50° angle, numerous barely weak primary lateral straight or weakly curved to the collective veins (barely distinguishable from the veins, weakly sunken above and about as interprimary veins) rather than 8-10 distin prominent as collective veins, raised below guishable primary lateral veins. All the and more prominent than primary lateral lateral veins of A. chocoense are also more veins, drying prominently raised on both prominently raised upon drying than the surfaces; interprimary veins numerous, veins of A. rupicola. drying prominulous on both surfaces; The species was first collected in 1955 by collective veins drying raised on both Jesus Idrobo. All collections are known surfaces, generally more prominent than from the Department of Choco, hence the the lateral veins, not loop-connecting the name A. chocoense. primary lateral veins. INFLORESCENCE erect-spreading, usually several per plant Additional specimens seen-COLOM at flowering; peduncle terete, 30-53.5 cm BIA. Choco: Condoto vic., Idrobo 1901 long, 2-3 mm diam.; spathe green, sub (COL); Mun. Quibdo; Corr. de Bebara, coriaceous, narrowly lanceolate, 3.5-4 cm sector La Calle, in Rio Bebara, Cordoba X 5-8 mm, broadest usually just above the 331 (MO); 441 (CHOCO, COL); 492 (COL); base, abruptly to gradually acuminate at Corr. de San Francisco de leho, Rio apex, acute at base, inserted at 45° angle on Necoram, Galeano & Bernal 429 (HUA); peduncle; spadix white at anthesis, be Correg. Guayabal, Rio Hugon, Forero & coming green post-anthesis, weakly ta Jaramillo 2791 (COL). pered to apex, 3.7-6.0 cm X 3-6 mm near the base, 2-3 mm diam. near the apex. Anthurium palacioanum Croat, sp. nov. Flowers subquadrangular, 2.0-2.2 X 1.4- Type: ECUADOR. Napo: Canton 1.6 mm, 5-6 flowers visible in the principal El Chaco. RIO Granadillo, Campa spiral; tepals semiglossy, the lateral tepals mento de INECEL, "Codo Alto", 2-2.2 mm wide, the inner margin thin, 0008'S, 7r28'W, 1300 m, W Palacios T. B. CHOAT, .J. WHITEHILL, E. D. YATES, 2007 33

Fig. 4. A. Antburi/lln vanderknaajJii Croa t. (Croa / 9034]). Inflo rescence , close-up. B. A n/huriLim we/.ffii Croa t. (van der \'(/el.//e/ at. 1l98Z). Herbariu111 type speci111 en.

5580 ( ho lotype, MO-4372260; iso fu sed 5-7 111111 to the adjacent pair of basa l types COL, K , QCNE, US). Figure 3B ve ins; primary lateral veins 4 pairs per side, arising at 25-30° angle; collecti ve Planta rupicola; internodia brevia, 1 cm veins ari sing fro m the 1st basa l vein, diam.; ca taphylla 5.5 C111 lo nga, non persis s0 111 etimes weakly loop-connected w ith tentia; petio lus 11- 19 cm lo ngus, teres ; th e 2nd ba sa l vein. INFLORESCENCE held lamina 13.5-18 C111 lo nga, 9.5-12 cm lata, slightly above th e leaves; peduncle robust, nervi pri111a rii laterales 4 utroque; pedun 31 cm lo ng; spathe green, oblong-Ianceo culus 31 C111 lo ngus; spatha oblongo-Ian late, erect-sprea ding to retlexecl , 4.3 C111 ceolata, 4.3 cm longa, 1 cm lata . long, ca . 1 cm \v ide, narrowly acuminate, Rupicolo us; ste111 s sho rt; internodes inserted at an acute angle; spadix 4.7 C111 sho rt, ca . 1 C111 cliam. ; roots moderately long, stipe 1.5 111111 lo ng, spadix proper few, drying 1.5 111111 diam.; cataphylls to 3.2 C111 long, drying 6 111111 dia111. , w hite to 5.5 cm long, light reddish brown, intact crea 111 , tapered to a blunt apex. Flo w ers ca . w ith pale fi bers at base, apparently de 4 per spiral, 1.4-1.7 X 2.0-2.2 m111 ; tepals ciduo us; LEAYES erect to spreading; pe w ith a few raphide cells; lateral tepa ls tioles 16-19 cm long, d rying ca. 3 mm 1.6 mill w ide, the inner Illargin broadly d ia111. , terete, slightly lo nger than the rounded. INFRUCTESCENCE not seen. blades; geniculum 1. 5 cm lo ng, as th ick as A n/burium jJalacioanum is ende111i c to petio les ; blades subcori aceous, epunctate, Ecuador, knmvn o nly fro m two localiti es in ovate-rhombic, 13.5-18 X 9.5-12 cm, grad Na po Province, o ne at the type locality in ua lly acu111inate at apex, broa d ly obtuse to Ca nton EI Chaco, the other in Canton almost truncate at base, d rying gree nish to Archidona o n the H o llIn-Lo reto Roa d. The gra yish yellow-brown above, gray-gree n to species usually occurs o n rocks in rapidly pa le yellow-brown below; lnidrib and tl owing strea111 S at 800-1980 111 elevation in basa l veins moderately raised o n drying; Premon/ane rain forest (P-rf). The species ba sa l veins 3-4 (-5) pa irs, all free to the is named in honor Walter Pa lacios, former ba se o r sometimes 'w ith th e lowermost ly an employee o f the H erbario Nacio nal 34 AROIDEANA, Vol. 30 and one of the finest plant collectors in the erect, becoming prominently reflexed, nar history of collecting in Ecuador. Walter has rowly acuminate at apex (the acumen 1- collected many interesting plants and new 6 mm long, inrolled), inserted at ca. 45° species, including this species. angle; spadix stipitate 2-3 mm, greenish brown (probably at anthesis), narrowly Additional specimens seen-ECUADOR. oblong, scarcely or not at all tapered, 7.4- Napo: Codo Alto, RIo Granadillo,faramillo 9.5 cm X 3 mm (dried), bluntly rounded at et at. 12681 (COL, HUA, MO); Cant6n apex. Flowers rhombiC, 2.2-2.5 X 1.9- Archidona, Hollfn-Loreto Rd., RIO Ruatar 2.2 mm, the sides gradually sigmoid; 3 aco, 2 hrs. by foot to Aldea Guagua flowers visible in the principal spiral, 5-6 Sumaco, Ceron & Factos 7441 (MO). flowers visible in the alternate spiral; tepals drying glossy, lateral tepals 1.3-1.4 mm Anthurium sytsmae Croat, Monogr. Syst. wide, inner margins convex, thin; pistils Bot. Missouri Bot. Gard. 14: 185-186. not at all emergent, green; stigma slit-like, 1986. Type: PANAMA. Code: area 0.3 mm long, the space between the tepals between Cano Blanco del Norte, Cano 0.3-0.4 mm wide; staminal emergence not Sucio and Chorro del Rio Tife, 8.42'19/1- studied, stamens apparently retracting (at 8.43'06/1N, 80.36'30/l-80.38'W, 200- least upon drying); anthers 0.3 mm long; 400 m, evergreen forest, 3 Feb. 1983, thecae oblong-ovoid, slightly divaricate. G. Davidse & c. W Hamilton 23578 INFRUCTESCENCE spreading; spathe per (holotype, MO-3099553; isotypes, K, sistent; berries green, obovoid, round at PMA). Figure 3C. apex, 6.5-9 X 4.5-5.5 mm, periderm thick, lacking obvious raphide cells; seeds 2, 4.5- Rupicolous; 35-50 cm tall; stems ca. 5 X 2-2.8 mm, somewhat flattened, envel 10 cm long; roots moderately few, des oped in a gelatinous substance. cending, ca. 3 mm diam.; cataphylls 6- Antburium sytsmae is endemic to Pana 9 cm long, narrowly pointed at apex, thin, ma in the Cocle and Bocas del Toro soon weathering to slender, pale brown Provinces occurring in Tropical wet forest fibers and persisting loosely at uppermost (T-wt). It is known only from rapidly nodes, eventually falling. LEA YES erect to flowing streams at 200-400 (850-950) m spreading; petioles 13-18 cm X 2-3 mm, where it grows on rocks along the stream narrowly sulcate; geniculum 1.3-1.5 cm margins or in the spray basin of waterfalls. long; blades subcoriaceous, oblong-oblan The species is closely related to A. ceolate, 19-25 X 3-4.5 cm, gradually acu amnicola, which is also only sparsely and minate at apex (the acumen 1-2 cm long, obscurely punctate on the upper surface, as apiculate ca. 1 mm), narrowly acute at base, well as A. antioquiense Engl., from Colom epunctate below, drying yellowish brown, bia, which is only sparsely punctate. sparsely and obscurely glandular-punctate This species was the third riparian above, drying grayish brown; midrib species to be discovered in rapidly flowing raised below, weakly raised above; prima streams on the Caribbean slope of Cocle ry lateral veins 5-7 pairs per side, Province (the other two being Antburium prominently ascending, departing midrib rupicola and A. amnicola). It differs from at 15-30° angle, drying scarcely more both of these species in that it has prominent than the interprimary veins; prominent reticulate venation. Although reticulate veins drying prominulous on this venation is closest to that of A. upper surface, prominent on lower surface; am nicola, the latter species has an ovate, collective vein arising from one of the pale lavender spathe and a much shorter, primary lateral veins in the middle to lower violet-purple spadix. It differs from A. third of the blade. INFLORESCENCE erect, rupicola because the collective vein arises usually longer than the petioles; pedun from one of the primary lateral veins in the cles 13-16 cm long; spathe oblong-lance middle of the blade. Antburium rupicola olate, 2.7-4.7 cm X 5-10 mm, green, at first has the collective vein arising from the base T. B. CROAT, J. WHITEHILL, E. D. YATES, 2007 35 of the blade. Anthurium sytsmae also lacks and obscurely densely short-lineate, drying the leaf punctations that are clearly present light to medium reddish brown; geniculum in A. rnpicola. 5-15 mm long, scarcely thicker, obtusely C Anthurium sytsmae has been collected shaped to sharply C-shaped, blades sub in flower and fruit in February and is coriaeeous, dark green and moderately named for Kenneth Sytsma who made the glossy above, slightly paler to slightly less first collection in 1982. glossy below, drying dark green to olive green above, slightly paler yellow-green Additional specimens seen-PANAMA. below, laneeolate, (8-)13-14(-19) em long, Cocle: Davidse & Hamilton 23639 (MO); (2.8-)3-4.5 (-6.3) em wide at middle; mid vic. of waterfall of Rio Tife, base of Cerro rib raised and paler (drying coneolorous) Tife, Knapp 3699 (MO, US); between Cafia above, drying narrowly raised and slightly Susio and waterfall at base of Cerro Tife, ca. paler yellow below; primary lateral veins 4 hr. hike, Sytsma 2544. Bocas del Toro: ca. 5-7 pairs per side, weakly etched above, Gualaca-Chiriqui Grande, along the oil slightly darker and flat below. INFLORES pipeline N of Continental Divide, in forest CENCE erect; peduncle 14-17.5 em long, W of pipeline rd. at end of driveable rd., 1.5-2 mm diam., drying dark reddish Hammel et al. 14703 (B, MO, PMA). brown, slightly ribbed; spathe broadly ovate, 4-4.5 em long, 2.5-3.8 em wide at Anthurium vanderknaapii Croat, sp. widest point, almost as broad as long, nov. Type: COLOMBIA. Valle: Dagua, subeordate, emerging white, soon becom originally collected by Arend de ing red-orange, then greenish in age, drying Willde, 1996, cultivated at Anthura, reddish brown, weakly glossly above, Inc., Bleiswijk, Holland, vouehered semiglossy below; spadix bright golden Mar. 2003, as T B. Croat 90341 yellow to orange, 3.5-6.5 em long, 4-6 mm (holotype, MO-5864048; isotypes, B, wide before anthesis, up to 1.7 em wide at CAS, COL, GB, K, NY, US). Figur anthesis, ageing to a dirty pink, drying dark es 3D, 4A. brown to dark reddish brown. Flowers 5-6 per spiral, square, the margins straight on Terrestris, rupicola, ad 1.5 m; internodia the principal spiral, weakly sinuate on the 2-6 (-7) em longa, 5-8 mm diam.; eata perpendicular margins; tepals weakly phylla 5-8.5 em longa, persistens semi glossy; pollen yellow; berries orange-red; intaetum; petiolus 4-9 (-13.5) em longus, seeds 2.25 mm long, 1.5 mm diam.; meso ca. 2 mm diam.; lamina laneeolata, (8)13- carp juicy. 14(19) em longa, (2.8)3-4.5 (-6.3) em lata; Anthurium vanderknaapii is endemic to nervi primarii laterales ca. 5-7 per utroque; Colombia, known only from two localities, pedunculus 14-17.5 em longus, 1.5-2 mm the type locality on the western slope of the diam.; spatha late ovata, 4-4.5 em longa, Cordillera Occidentale, as well as near 2.5-3.8 em lata, subcordata, rubra-auran Manizales in the Cordillera Central. The tiaeus: spadix aureus vel aurantiacus, 3.5- species is recognized by its elongate inter 6.5 cm long, 4-6 mm diam. anthesis, ad nodes, lanceolate, eglandular blades, 1.7 em diam. per anthesis. broadly ovate red-orange spathe, and Terrestrial, rupicolous, 1.5 m tall; inter golden-yellow spadix. No other species of nodes 2-6 (-7) em long, to 5-8 mm diam., Anthurium appears to be confused with medium to dark green, semiglossy; roots this species and none have this combina a few at each node; cataphylls 5-8.5 em tion of characters. It is placed within long, turning reddish brown, sharply 1- section Calomystrium because of its color ribbed near apex, persisting semi-intact at ful spathe and spadix, characteristic of this upper nodes; petioles 4-9 (-13.5) em long, section, and placed within the subsection ca. 2 mm diam., subterete to weakly and Rupicola due to its original collection near narrowly flattened, weakly sulcate adaxi Dagua in an area of many streams and rock ally, medium dark green and semiglossy outcrops. 36 AROIDEANA, Vol. 30

This new species is named for Nic van 1.5 cm long, drying darker than petiole; der Knaap, owner of Anthura, Inc., Bleis blades narrowly oblong-elliptic, reticulate wijk, Holland, where the type plant was ly veined, epunctate, about the same length vouchered by the senior author. Anthura, as the petioles, subcoriaceous, oblong Inc. is the world's largest breeder of oblanceolate, 15-23 X 2-3.5 cm, gradually ornamental Anthurium, and has contribut acuminate at apex (acumen 1-1.5 cm long, ed immeasurably to the worldwide interest apiculate 2 mm), narrowly acute at base, and popularity in ornamental aroids. Nic drying yellowish green; midrib weakly van der Knaap has been benefactor of the raised above, round raised below; prima International Aroid Society being one of ry lateral veins 5-7 pairs per side, only a few financial sponsors of the VIII prominently ascending, departing midrib International Aroid Conference in St. Louis. at 20-30° angle, drying yellow below (otherwise tertiary veins nearly as promi Additional specimens seen-COLOM nent); basal veins 1 pair, rising from the BIA. Caldas: Manizales, collected and base, collective vein arising from primary cultivated by Marta Posada de Robledos, lateral veins in the middle to lower third of Medellfn, Colombia, vouchered 28 Apr. the blade; INFLORESCENCE erect, much 1983 as Croat 56358 (MO). shorter to slightly longer than petioles 02- 25 cm); peduncles 8-22 cm long, slender; Anthurium wer:lfii Croat, sp. nov. Type: spathe oblong-Ianceolate, 2-2.5 cm X 6- ECUADOR. Esmeraldas: border be 8 mm, spreading to reflexed, acute to tween Carchi and Esmeraldas, ca. acuminate at apex, green tinged with 20 km past Lita on road Lita-Alto purple, inserted at ca. 45° angle; spadix Tambo, 0045'4"N, 78°15'4"W, 550 m, stipitate (stipe 2-3 mm long), spadix yel H. van der Wer:lJ; B. Gray & G. Tipas low or green, cylindric to slightly tapered 11981 (holotype, MO-3870548; iso toward apex, 1.5-3.5 cm X 3 mm at types, B, CAS, COL, F, K, NY, QCA, anthesis. Flowers square, 1.7-2.0 X 1.8- QCNE, US). Figure 4B. 2.0 mm, the sides perpendicular to spirals, Planta epilithica; internodia brevia, 1 cm Sigmoid; 3 or 4 flowers visible in both diam.; cataphylla 5-7 cm longa, persisten principal and alternate spiral; lateral tepals tia in fibras tenues soluta; petiolus 11- 1.0 X 0.7 mm, inner margins convex; pistils 23 cm longus, teres aut leviter sulcatus; not emergent. INFRUCTESCENCE erect; lamina subcoriacea, oblongo-oblanceolata, spathe persistent; berries globose, round 15-23 cm longa, 2-3.5 cm lata, epunctata; at apex, 4.7 X 3 mm, each purple on upper nervi primarii laterales 5-7 utroque; nervi third, without visible raphide cells, bilocu collectivi e nervis primariis lateralibus lar; seeds one per lonile, 3 X 1.6 mm, medio vel triente inferiore laminae exor 1.4 mm thick, with small transparent muci ientes; inflorescentia erecta; pedunculus 8- lage at micropylar end. 22 cm longus; spatha viridis, oblongo Anthuriu111 werjfi is known only from lanceolata, 2-2.5 cm longa, 6-8 m111 lata; the Pacific slope of Ecuador in Esmeraldas spadix viridis, stipitata 2-3 mm, 1.5-3.5 cm Province at (200) 500-550 m in Tropical longa, 3 mm diam.; baccae purpureae. rainforest (T-rt) and Tropical wetforest (T Epilithic; stems ca. 5 cm long; inter wf). This species grows principally on nodes short, less than 1 cm diam.; roots rocks or trees along and within streams. moderately few, ca. 3 mm diam.; cata The species is named for Henk van der phylls 5-7 cm long, soon weathering into Werff, curator at the Missouri Botanical thin, yellow-brown, persistent fibers, 1-2 of Garden, who made the first collection of the uppermost persisting intact, dark the species and the type specimens. Henk, brown. LEA YES erect, soon spreading to harking back to his Dutch background, has reflexed; petioles 11-23 cm X 2 mm, an interest in bringing interesting plants terete or slightly sulcate; geniculum 1- into cultivation. T. B. CROAT, J. WHITEHILL, E. D. YATES, 2007 37

Additional specimens seen-ECUADOR. rupicolous species seem to have evolved Esmeraldas: Rio Negro, 2 km from El blade shapes that can withstand the Placer, Palacios 6921 (CAS, MO, QCNE); tremendous shearing pressures of rushing San Lorenzo Canton, Res. Etnica Awa, water in those relatively brief periods Parroquia Ricaurte, Centro Pambilar, Au when streams rise following heavy rain lestia & Aulestia 1058 (CAS, MO, QCNE); storms. It is curious that A. rupicola is the Parroquia Mataje, Centro Mataje, Aulestia et only species in section Porphyrochitonium al. 488 (MO); up Rio Palavi from Awa occurring on rocks that has made the Indian encampment, Hoover et al. 4115 transition into streams. Other species in (MO, QCA); 4167 (MO, QCA); 4176 (MO, the section Porphyrochitonium occur as QCA); Res. Cotacachi-Cayapas, Rio Tigre, epiphytes or are terrestrial understory Comejo & Bonifaz 6499 (GUAY, MO). forest species. Since Anthurium section Calomystrium provides the bulk of the CONCLUSION species used in the cut flower industry These nine moderately to highly en these lanceolate species of Calomystrium demic Anthurium species occur as rupi may be impOltant members of breeding colous stream bed epiliths. In an earlier programs. Already A. amnicola has played treatment, the senior author assigned three a very important role in providing a laven of these species to section Porphyrochito der-colored spathe in hybrids. nium on account of their punctate leaves (Croat, 1986). Although leaf punctations liTERATURE CITED are clearly present in A. rupicola they are T. quite sparse and inconspicuous in A. Croat, B. 1986. A revision of the genus amnicola and A. antioquiense, and, on Anthurium (Araceae) of Mexico and second examination of A. sytsmae, appear Central. America. Part II: Panama. entirely absent in that species. Leaf punc Monogr. Syst. Bot. Missouri Bot. Card. tations are also absent from A. antro 14:1-204. phyoides, A. palacioanum, and from one --- & G. S. Bunting. 1979. Standardi of the new species, A. werffii. There is zation of Anthurium descriptions. now evidence from breeding programs Aroideana 2:15-25. with some species in this complex, espe Dressler, R. L. 1980. A new name for the cially A. amnicola, that all of these species dwarf purple Anthurium. Aroideana are actually members of section Calomy 3(2):55. strium. Anthurium amnicola will not Engler, A. 1905. Araceae-Pothoideae. pp. 1- cross with members of section Porphyr 330 In A. Engler (ed.), Das Pjlanzen ochitonium, but it does cross readily with reich IV 23B CHeft 21). W. Engelmann, members of section Calomystriul11. Many Leipzig and Berlin. typical members of section Calomystrium Holdridge, L. R., W. H. Hatheway, T. Liang have glandular punctations, but until now & J. A. Tosi. 1971. Forest Environ no Ian ceo late-bladed species were consid ments in Tropical Life Zones. Perga ered members of this section. These mon Press, New York.