Contributions from the United States National Herbarium

Volume 35


Ellsworth P. Killip, Jose Cuatrecasas, Lyman B. Smith, and Joseph Ewan


Publications of the United States National Herbarium

The United States National Herbarium, which was founded by the Smithsonian Institution, was transferred in the year 1868 to the Department of Agriculture and continued to be maintained by that department until July 1, 1896, when it was returned to the official custody of the Smithsonian Institution. The Department of Agri- culture, however, continued to publish the series of botanical reports entitled "Contributions from the United States National Herbarium," which it had begun in the year 1890, until, on July 1, 1902, the Na- tional Museum, in pursuance of an act of Congress, assumed responsibility for the publication. The first seven volumes of the series were issued by the Department of Agriculture.

Frank A. Taylor, Director, United States National Museum.



Part 1, Supplemental notes on the American species of Passifloraceae, with descriptions of new species. By Ellsworth P. Killip. (Published May 10, 1960) 1-23

Part 2. A taxonomic revision of the Humiriaceae. By Jose

Cuatrecasas. (Published April 14, 1961) .... 25-214

Part 3. Origins of the flora of southern Brazil. By Lyman

B. Smith. (Published March 19, 1962) .... 215-249

Part 4. A Synopsis of the American Velloziaceae. By Lyman

B. Smith. (Published March 19, 1962) .... 251-292

Part 5. Synopsis of the South American species of Vismia (Guttiferae). By Joseph Ewan. (Published June 5, 1962) 293-377

Part 6. Cacao and its allies: A taxonomic revision of the genus Theobroma. By Jose Cuatrecasas. (Pub- lished August 21, 1964) 379-614



Contributions from the United States National Herbarium

Volume 35, Part 1




By Ellsworth P. Killip

Bulletin of the United States National Museum SMITHSONIAN INSTITUTION * WASHINGTON, D.C. 1960


This paper was partly prepared by Ellsworth P. Killip, 10 years after the publication of his monograph of the Passifloraceae in 1938, to summarize the additional data acquired in the 10-year period, and to present descriptions of 8 new species. In 1956, Mr. Killip turned over the manuscript to Dr. Jose Cuatrecasas to use as he saw fit in the preparation of a flora of Colombia. It is now published with the addition of three new species that Mr. Killip indicated as new but did not describe. Dr. Cuatrecasas wrote descriptions for these as well as Latin diagnoses for all the new species described by Mr. Killip. Included are 11 new species and citations of all new material examined since the publication of the monograph. Dr. Lyman B. Smith, curator of the division of phanerogams, has care- fully edited the manuscript.

Jason R. Swallen

Head Curator, Department oj Botany




By Ellsworth P. Killip


In 1938 the Chicago Natural History Museum (then the Field Museum of Natural History) published a monograph on the American species of Passifloraceae written by the present author.1 The prepara- tion of this monograph covered a period of many years, during which time I made three visits to Europe and consulted nearly all the type specimens as well as many others of historic importance. A large amount of material was generously placed at my disposal by those in charge of many of the larger herbaria of the world. It is therefore perhaps not surprising that in the intervening years I found little reason to revise the treatment of the species presented in the mono- graph.

The intensive exploration carried on in certain parts of northern South America, especially in the basins of the Orinoco and Amazon in Colombia and Venezuela, resulted in the collecting of many Passi- floraceae in regions little known in 1938. In the course of studying these specimens, as well as material from Central America and other parts of South America, I detected several new species, some of which I described elsewhere. This material also shows interesting extensions of ranges of certain species, and in some instances permits a desirable amplification of the descriptions given in the monograph. It therefore seems advantageous to assemble in one place all this subsequent information along with descriptions of several new species. The numbers associated with the species of Passifiora are those of the monograph.

4. Passifiora gracillima Killip

This species is now represented in the U.S. National Herbarium by several additional Colombian collections. One specimen from the

> Field Mus. Bot. 19. 1938.


Province of Chiriqui, Panama, was cited; the species was recollected in the vicinity of Chiriqui Lagoon in 1941 by H. von Wedel (No. 2421). Likewise, a single specimen was listed from Riobamba, Ecuador. This species is now represented by Steyermark 52932, which was collected between Cruz Pamba and Loma de Canela, north of Molieturo, Province of Azuay, Ecuador, at an altitude of 2,315-2,500 m. The collector noted that the fruit is edible and has a flavor similar to that of Annona.

10. Passiflora gracilis Jacq.

Additional collections are:

Venezuela: Bolivar: Gran Sabana, Cerro Kanaime Santa Elena, March

1946, Tamayo 3025. Ecuador: Azuay: Between Rio Gamolotal and Rio Norcay, Steyermark 52879. Both of these collectors noted that the fruit is red. Steyermark gave "shunshun" as a local name, and stated that the corona fila- ments are white with a purple streak at the center and that the seeds are red.

14a. Passiflora ursina Killip & Cuatr., sp. no v. Plate 1

Frutex elatus scandens; rami foliosi dense rufo-tomentosi; stipulae subulatae; folia coriacea trinervia ovato-oblonga vel oblonga basi truncata vel rotundata apice acuta vel subacuta mucronulata utroque latere saepe 1-lobata, lobo triangulari brevi mucronulato, subtus rufo-hirtella vel tomentella, margine integra revolutaque, petiolo brevi basi duobus glandulis minutis inconspicuis; bracteae liberae subulatae; calycis tubus petalliformis, sepalis ovatis apice rotundatis subapicem corniculatis, petalis paulo brevioribus; corona filamentis biseriatis apice incrassatis cxterioribus robustioribus; operculum membranaceum plicatum erectum breviter fimbriatum; ovarium glabrum.

Large, scandent shrub; leafy branches moderately thin, flexuose subterete, densely reddish brown tomentose with acute, subrigid, spreading hairs, 0.5-1 mm. long; tendrils closely spiral, tomentose; stipules subulate, hirsute, 5-6 mm. long, 0.5-1 mm. broad; petioles 8-14 mm. long, stout, straight or bent at base, densely rufo-tomentose (like the branches), the glands small, semiglobular, sessile at base, concealed by the indument; blades coriaceous, trinerved, ovate-oblong or oblong, 5-8.2 cm. long, 3-4.5 cm. broad, rounded, truncate or cordate at base, subtrilobate (or rarely entire) near the apex, the two lateral lobes very short, form'ng an obtuse mucronulate angle, the terminal one (the apex) acute or subacute, mucronulate, the margin entire, revolute, the surface puberulous with spreading, sparse hairs


above, pubescent or subtomentose with spreading, long, rufous hairs beneath, the 3 nerves and veins very prominent beneath, little con- spicuous above; a few minute, semiglobose glands near the base beneath; inflorescence cymose, the cymes in pairs with 5-7 flowers, the rachis less than 1 cm. long; bracts subulate, 3-5 mm. long, up to 0.5 mm. broad, rufo-hirsute; the peduncles thick, straight, rufo- tomentose, 8-14 mm. long, articulate about the middle or slightly above the middle; flowers 2.5-3 cm. wide, greenish white; sepals ovate, obtuse, 1.1-1.3 cm. long, 0.7-0.9 cm. wide, dorsally more or less hirsute and keeled, corniculate below the apex, the horn up to 3 mm., hairy; petals membranaceous, ovate, narrowed at both ends, obtuse, subdentate below the apex, 1-1.1 cm. long, 0.6-0.7 cm. wide; corona filaments filiform, much dilated at apex, in 2 series, the outer about 3.5 mm. long, rigidulous, the inner about 2.5 mm. long, thinner and flexuose; operculum membranaceous, closely plicate, short- fimbrillate, 1.5 mm. high; limen annular; gynophore absent; ovary ellipsoid-obovoid, glabrous.

Type in the U.S. National Herbarium, No. 1799118, collected at Paramo de Santa Lucia, south side of Laguna de la Cocha, Depart- ment of Narino, Colombia, at an altitude of 2,900-3,100 m., on Jan. 9, 1941, by J. Cuatrecasas (No. 11886).

P. ursina is closely related to P. apoda Harms and P. sodiroi Harms, from which it differs in the thick, longer, rufous indument, in the shorter petioles with sessile, minute basal glands concealed by the indument, and more conspicuously in the horned sepals. It differs from P. sodiroi in lacking the gynophore and in the form of the petals, and from P. apoda by the shorter corona filaments.

26. Passiflora dolichocarpa Killip

Known only from the type specimen at the time of my monograph, this species is now represented in the U.S. National Herbarium by the following additional collections:

Mexico: Chiapas: Hacienda Siltcpec, Matuda 1892.

Guatemala: Jalapa: Between Jalapa and Montana Miramundo, alt. 1,500-

2,000 m., Steyermark 32884 (F). Suchitep6quez: Volcan Santa Clara, alt.

1,250-2,650 m., Steyermark 46761.

46a. Passiflora maerostemma Killip, Field Mus. Bot. 23:217. 1947. Additional collections are:

Guatemala: Chiquimula: Volcan de Quezaltepeque, alt. 1,500-2,000 m., Steyermark 31519 (F, type, US photo and fragment), 31281 (F). Jalapa: Montana Durazno, alt. 1,400-1,900 m., Steyermark 32985 (F).


61. Passiflora lancearia Mast.

Recorded as being confined to Costa Rica, this species was collected in the Province of Chiriqui, Panama, in 1938 by M. E. Davidson (No. 336) and in the Province of Bocas del Toro in 1941 by H. von Wedel (No. 2120).

63. Passiflora panamensis Killip

This species was known at the time of my monograph only from the type, which came from southern Darien, Panama. The species has been found in the adjacent coastal part of Colombia, a recent record being:

Colombia: El Choc6: Bahia Solano, dense forest along the Quebrada Jellita, alt. 50-100 m., Feb. 22, 1939, Killip & Garcia 33482.

69. Passiflora bauhinifolia H.B.K.

P. andreana Mast, was treated as a synonym of P. bauhinifolia in my monograph. Specimens collected by Garcia-Barriga and Balls in southwestern Colombia in 1939 show that these two species are unquestionably distinct. In P. andreana the inside of the sepals and the petals are purple, reddish purple, or deep violet; in P. bauhinifolia these parts are white. The outer corona filaments are slenderer in P. andreana.

Of the specimens listed in my monograph under P. bauhinifolia, I have rechecked only those in the U.S. National Herbarium; on this basis, P. bauhinifolia is common in Ecuador, and P. andreana is known only from Colombia. P. bauhinifolia is represented in the U.S. National Herbarium by the following:

Colombia: El Valle: Quebrada de la Palma, Rio Bugalagrande Valley,

Cuatrecasas 20935. Putumayo: Laguna de la Cocha, Paramo El Tabano,

Garcia-Barriga 7785A. Ecuador: Without locality, Milk, Imbabura: Otovalo, Alexia 7645.

Pichincha: Quito, Humbold & Bonpland (photo of type); Mille 222, s.n.

in 1918. Verde Cruz, Firmin 310. Guapulo, Stevens 258. Tungurahua:

Banos, Lehmann, 675, 8020. Peru: Cajamarca: Pena Rota, toward Cochabamba 2,800 m., Ochoa 1565.

This last record is apparently the second of this species in Peru.

69a. Passiflora andreana Mast. Journ. Linn. Soc. 20:37. 1883.

Additional collections are:

Colombia: Narifio: Las Lajas, near Ipiales, Andr6 3478 (photo of type); Garcia-Barriga 7757A; Balls 7360. Jenoy, 5 km. northwest of Pasto, alt. 2,460 m., Ewan 15899 (USNA). Between Pasto and Anganoy, Schultes & Villarreal 7417.


100. Passiflora micropetala Mast.

In the monograph I stated that this species was one of the few maintained as valid by Masters in the Flora Brasiliensis that I knew only from description. After studying a specimen subsequently col- lected along the Rio Putumayo, I have decided that it and a speci- men from the Comisaria of Caquetd, Perez-Arbeldez 669, which was cited by me as P. dawei, both represent P. micropetala. The type locality of this species as given by Masters is : "In regione fluvii Japura, prov. do Alto Amazonas." The Rio Japura is identical with the Rio Caqueta, and the Putumayo is in the same general area. On his trip to this river, Martius is known to have collected in present-day Colombia, and the type may thus have come from this country.

The Perez collection is sterile, and the more recent one is in fruit only. Both resemble P. biflora in leaf shape, but the coronal structure is different according to Masters' description. P. micropetala there- fore appears to be known from the following locations :

Colombia: CaquetA: Florencia, alt. 420 m., Pirez-Arbelaez 669 (COL, US).

Putumayo: Selva higr<5fila del Rfo Putumayo en Puerto Ospina, alt. 230 m.,

Cuatrecasas 10776 (COL, US). Brazil or Colombia: Rio Japura (R. Caqueta*), Martius (M, type).

111. Passiflora adenopoda DC.

A recent collection of this species by Dugand and Jaramillo (No. 3965) has far more deeply cleft bracts than heretofore observed. They are deeply palmate-trisect, and the segments are repeatedly cleft. The largest bract is 2 cm. wide. This unusual variant was collected in the Department of Cundinamarca, Colombia, at Es- taci<5n San Bernardo, between Sasaima and Alb&n, at an altitude of 1,700-1,800 m.

132. Passiflora bicuspidata (Karst.) Mast.

This species, known hitherto only from the Eastern Cordillera of Colombia, was recently collected by Steyermark across the border in Venezuela. His detailed notes on the color of the flowers are worth reproducing as earlier collectors have described the flower color in only general terms.

"Calyx rose-lavender, the lobes with more lavender, the tube with more brick-color; coronal filaments golden; catyx tube below the coro- nal filaments within dull pale greenish-yellow; filaments purple; an- thers buff; styles purplish; petals purplish-rose."

The collection was made at:

Venezuela: Tdchira: Swampy woods between Paramito and Quebrada de Palmar, at base of Pdramo de Tamd, 7 km. above Villapaez, alt. 2,500 mv Steyermark 57240 (F, US).


135. Passiflora sanguinolenta Mast.

In the monograph I followed Masters in placing this Ecuadoran species in the subgenus Psilanthus, but I observed that it appeared more closely related to P. rubra, of the small section Xerogona of the subgenus Plectostemma, than to the three known species of Psilanthus.

Later, there were submitted to me, through the courtesy of Harold N. Moldenke, then of the New York Botanical Garden, a fine series of specimens of P. sanguinolenta and P. rubra collected by Reinaldo Espinosa in the Province of Loja. One of P. sanguinolenta, No. 806, is in fruit, apparently the first time that fruit of this species has been collected. The transverse ridges of the seeds are rugose, not smooth as in Xerogona, and I believe that the species is best left in Psilanthus, even though in vegetative parts (especially in the absence of any bracts) and shape of the fruit it very closely resembles P. rubra.

To the collections cited in the monograph there may now be added the following, all from the Province of Loja and deposited in the U.S. National Herbarium:

Ecuador: Catacocha, alt. 2,050 m., Espinosa 617. Pucala, north of Loja, alt. 2,300 m., Espinosa 621. Between San Pedro and Chinchas, about 55 km. west of Loja, alt. 1,200 m., Espinosa 1314. Aguahedionda, north of Loja, alt. 2,150 m., Espinosa 806. Desert country between Vilcabamba and Cachiyaeu, alt. 1,600-2,100 m., Steyermark 54397.

137. Passiflora bracteosa Planch. & Linden

Tacsonia infundibularis Mast.

Both these species wore based upon Funck and Schlim's 1381. The type locality of P. bracteosa was correctly given as La Baja, Colombia; that of Tacsonia infundibularis was incorrectly recorded as "Vene- zuela." However, this species has recently been collected by Steyer- mark (No. 57277, at F) in the State of Tachira, Venezuela, at the base of the Paramo de Tama, at an altitude of 2,285-2,430 m.

147. Passiflora adulterina L.f.

Although this species is fairly common in the vicinity of Bogota, fruiting material was not found, apparently, until 1939. Cuatrecasas 5136, collected between ElDelirio and Guadalupe (eastern slope of the Cordillera Oriental, between 2,900 and 3,220 m. altitude), bears mature fruit, which may be described as follows :

Fruit broadly ovoid, 7.5 cm. long, about 4.5 cm. in diameter, apparently reddish brown, mottled with greenish yellow; seeds obovate, about 9 mm. long, and 4.5 mm. wide, reticulate, dark brown, lustrous.


148a. Passiflora cuatrecasasii Killip, sp. no v. Plate 2

Suffrutex data scandens, caule primum Ianato denique glabrato; stipulae subulatae; folia coriacea ovato-lanceolata vel oblongo- lanceolata plus minusve serrata basi rotundata vel subcordata apice acuta vel subobtusa subtus lanata, petiolo sub apice 2-3 glandulis; bracteae liberae ovato-oblongae vel anguste oblongae acutae semi- Iatae; calycis tubus cylindricus glaber, sepalis tubo valde brevioribus oblongis obtusis, petalis scpala subaequalibus; corona annularis undulata; operculum deflexum denticulatum leviter recurvatum; ovarium tomentosum.

Vino; stem subangular, lanate when young, the indument at length evanescent; stipules subulate, up to 1.1 cm. long, glabrous, purplish black below middle, the upper part spongiose-thickened, light yellow- ish brown, coriaceous, acuminate; petioles up to 2 cm. long, bearing 2 or 3 glands near apex; leaves unlobed, ovate-lanceolate or oblong- lanceolate, 6-11 cm. long, 3-6.5 cm. wide, acute or subobtuse, rounded or cordulate at base, serrate or serrulate, penninerved (princi- pal lateral nerves about 9 to a side), coriaceous, dark green, lustrous, and glabrous above, densely lanate beneath, except sometimes on the nerves and veins; peduncles 7-10 cm. long, slender, evanescently lanate, articulate about 1 cm. below the apex; bracts distinct to the base, ovate-oblong to narrowly oblong, 3-4 cm. long, 0.8-3 cm. wide, acute, sessile, finely serrulate; calyx tube cylindric, 8-10 cm. long, 1-1.5 cm. wide, greenish at base, red or purplish above, glabrous; sepals oblong, 3.5-4.5 cm. long, 1-1.5 cm. wide, obtuse, dorsally awned just below the apex (awn about 1.5 mm. long), pinkish purple; petals subequal to the sepals; corona a single indulate ridge at the apex of the tube; operculum borne near the base of the tube, 4-7 mm. long, dependent, the margin slightly recurved and minutely dentic- ulate; ovary ovoid, densely tomentose.

Type in the U.S. National Herbarium, No. 1796435, collected on the eastern slope of the Paramo de Guasca, Cordillera Oriental, Depart- ment of Cundinamarca, Colombia, at an altitude of 3,200-3,300 m. on June 2, 1940, by J. Cuatrecasas (No. 9479). Also represented by J. Cuatrecasas 13538 from the same locality, and by Uribe 1061, from La Rusia, northwest of Duitama, Department of Boyaca, at an altitude of 2,700 m.

The proposed species probably should be placed nearest P. lanata in the arrangement adopted in my monograph, though it also has affinities with P. rugosa and P. leptomischa. From P. lanata it differs in having a scantier indument on the vegetative parts, a pubescent ovary, and generally longer peduncles. In P. rugosa the bracts are coalesced partway and the calyx tube is much shorter. P. leptomischa is a species with dimorphic foliage and very long (15-25 cm.) peduncles.

512425—60 2


148b. Passiflora rigidifolia Killip sp. nov. Plate 3

Caulis scandens angulatus sursum albopilosus basin versus gla- bratus; stipulae anguste lineares glanduloso-serrulatae subrevolutae; folia coriacea lanceolata integra basi rotundata apice acuta praeter costam supra pilosulam glabra, petiolo albo-piloso 2-3 glandulis parte media; bracteae liberae lanceolatae integrae; calycis tubus cylindricus glaber sepalis tubo brevioribus oblongis obtusis, petalis sepalis simi- libus; corona annularis dentata; operculum deflexum recurvatum minuteque denticulatum ; ovarium tomentosum.

Stem subangular, white-pilose toward the apex, becoming glabrous below; stipules narrowly linear about 1 cm. long and 1 mm. wide, acuminate, glandular-serrulate, subrevolute, coriaceous; petioles up to 7 mm. long, 2- or 3-glandular near the middle, white-pilose; leaves lanceolate, 4-6.5 cm. long, 1-1.8 cm. wide, acute at apex, rounded at base, entire, revolute (?), penninerved (midnerve impressed above, the lateral nerves and veins somewhat elevated on both surfaces), pilosulous on the midnerve above, otherwise glabrous, coriaceous, glaucescent beneath; peduncles solitary in the axils, 2-2.5 cm. long, glabrous or sparingly pilose; bracts borne about 5 mm. below the apex of the peduncle, free to base, lanceolate, about 2 cm. long and 8 mm. wide, acuminate, sessile, entire, pilose; flowers pale rose; calyx tube cylindric, 6.5-7 cm. long, 8-9 mm. in diameter, slightly enlarged at base, glabrous; sepals oblong, about 2.5 cm. long, and 1 cm. wide, obtuse, dorsally aristate just below apex, the awn 2 mm. long; petals similar and subequal to the sepals, about 8 mm. wide; corona a dark ring at the throat of the tube, etuberculate; operculum borne close to base of tube, 4 mm. long, dependent, the margin recurved and minutely denticulate; ovary ovoid, densely tomentulose.

Type in the Herbarium of the Royal Botanic Gardens, Kew, collected in Colombia at an altitude of about 3,900 m., 1894-1896, by D. Burke (No. 184).

This species is nearest to P. cuatrecasasii but differs in its much narrower leaves, shorter peduncles, and slenderer calyx tube.

Unfortunately, the locality at which the type was collected is uncertain. The label reads "Flora of Colombia. Locality: Conig, altitude 13,000 ft., Comm. Messrs. J. Veitch, 1897." The word "Conig" is not a Spanish one, and I have been unable to find in any gazeteer a Colombian name at all similar to it. Conejo, in Antioquia, on the railroad from Puerto Berrio to Medellin, suggests itself, but there is no altitude of 13,000 ft. attainable from there. In Peru there is a "Conec," but it is not known that Burke collected in that country.

Accompanying this specimen is a letter dated April 4, 1902, from James Veitch & Sons to Hemsley, in response to a request for infor- mation about Burke's collections. Nos. 146, 154, and 167 are listed


without localities, but the Veitch letter observes that the preceding numbers were collected in Antioquia. Nos. 184 and 185 follow both from "Conig," number 185 being given as "Berberis sp {Goudoti ?), 11,000 ft." Berberis goudoti is known only from the eastern Cordillera of Colombia.

149- Passiflora truxillensis Planch. & Linden.

This rare species hitherto known only from two collections from the State of Trujillo, Venezuela, was recently collected in the neighbor- ing State of Merida by Steyermark, from whose notes the following more complete description of the flowers may be presented:

Bracts pale green with pale rose line down the center; calyx tube pale green at very base, dull rose without, a greenish part in the middle; sepals deep rose at margin, lavender in the middle; petals rose-colored throughout; stamen-filaments rose-orchid; ovary green; styles rose-lavender above, whitish below.

The complete collection data are:

Venezuela: Merida: Rich forest between Mesa del Trapiche and El Molino, alt. 2,530-2,590 m., Steyermark 56503 (F, US).

154a. Passiflora boyacana Killip sp. nov. Plate 4

Herba scandens, caule lanato denique glabrescenti ; stipulae lineari- subulatae; folia coriacea profunde trilobata basi subtruncata vel sub- cordata lobis lineari-lanceolatis subacutis mucronulatis integris, subtus dense rufo-lanata, petiolo eglanduloso lanato; bracteae liberae integrae extus lanatae; calycis tubus cylindricus glaber, sepalis petalisque tubo dimidio brevioribus elliptico-oblongis obtusis mucronulatis; corona brevem annulum minute tuberculatum reducta; operculum deflexum recurvatum denticulatum; ovarium glabrum.

Herbaceous vine; stem stout, subterete, about 3.5 mm. in diameter, lanate, becoming glabrous; stipules linear-subulate, about 1.2 cm. long; petioles 1-1.5 cm. long, glandless, lanate in lines; leaf -blades lobed about three-quarters their length, 4-8 cm. along midnerve, 3.5-6.5 cm. along lateral nerves, 3-9 cm. between apices of lateral lobes, (lobes 1-1.5 cm. wide, narrowly lanceolate or the middle one sometimes elliptic-lanceolate, subacute, mucronulate), subtruncate or cordulate at the base, entire, coriaceous, essentially glabrous above with the nerves slightly impressed, densely rufo-lanate beneath; peduncles solitary, about 4 cm. long, pubescent like the petioles; bracts borne at base of flower, 2-2.3 cm. long, 8-10 mm. wide, free to the base, sessile, entire, deciduous-lanate without, glabrous within; flowers young; calyx tube cylindric, 4 cm. long, glabrous; sepals and petals equal, about 2 cm. long, obtuse, the sepals 7 mm. wide, mucron- ulate, the petals 1 cm. wide; corona reduced to a low, minutely tuber-


ciilate ridge; operculum borne at base of tube, dependent, about 4 mm. long, the margin recurved, denticulate; ovary ellipsoidal, glabrous.

Type in the U.S. National Herbarium, No. 1796049, collected in paramo thickets, above Puente de Boyaca, Department of Boyaca. Colombia, at an altitude of 2,800 m. on Feb. 24, 1940, by E. Perez- Arbelaez & J. Cuatrecasas (No. 8083).

This species, belonging to the subgenus Tacsonia, keys out to species 154-157 of my monograph of the American Passifloraceae. The narrow, entire leaf lobes are quite unlike those of P. ampullacea. In depth of lobation the leaves are rather suggestive of P. trianae and P. purdiei, which are essentially glabrous plants.

155a. Passiflora raimondii Killip, sp. nov. Plate 5

Herba scandens, ubique glabra; stipulae setaceae; folia membranacea trilobata, lobis oblongis vel ovato-oblongis, obtusis, denticulatis, peti- olis eglandulosis; bracteae liberae, lanceolatae, acutae; calycis tubus cylindricus, sepalis tubo brevioribus, lineari-oblongis, petalis anguste linearibus; corona annuliformis, crenulata; operculum deflexum.

Herbaceous vine, glabrous throughout; stem quadrangular, longi- tudinally ridged, the tendrils very slender; stipules setaceous, about 4 mm. long; petioles slender, 1-3 cm. long, glandless; leaves 3-5 cm. long, 4.5-8 cm. wide, 3-lobed to below the middle (lobes oblong or ovate-oblong, up to 3 cm. wide, rounded and sometimes slightly emarginate at apex, denticulate), cordate at base, membranous, bi- glandular at base; peduncles solitary, slender, about 5 cm. long; bracts lanceolate, 1-1.2 cm. long, 4-5 mm. wide at base, acute at apex, rounded at base, sessile, minutely denticulate, distinct to base, borne at base of flower; calyx tube cylindric, 3-3.5 cm. long, 7-8 mm. in diameter, narrowed at base; sepals linear-oblong, 2 cm. long, 3-3.5 mm. wide, obtuse, slightly carinate, the keel terminating in a filiform awn 1 mm. long; petals narrowly linear, 1.5 cm. long, 1.5 mm. wide, sub- acute; corona a low ridge, erenulate; operculum borne about 5 mm. above base of tube, dependent, 4 mm. long, minutely denticulate; gynophore very slender, about 5 cm. long; ovary fusiform.

Type in the Herbarium of the Universidad Nacional Mayor de San Marcos, Lima, collected at Chorrillos, Department of Cajamarca, Peru, at an altitude of 2,200 m. on June 8, 1875, by A. Raimondi (No. 7807). Represented also by A. Raimondi 8241, collected at Cascas, Province of Contumaza, in the same Department.

This species is most nearly related to P. gracilens, which is rather common in Peru and extends from Huanuco southward. The new species differs in its larger leaves with much broader lobes, its longer peduncles and larger flowers, and its- lanceolate, not ovate, bracts.


The Raimondi specimens were submitted to Harms, who suggested that they probably belonged to our undescribed species.

159a. Passiflora zamorana Killip sp. nov. Plate 6

Herba scandens folia supra et flores excepti dense griseo- vel brunneo- tomentosa; stipulae late ovato-reniformes semiamplexicaules plus minusve dentatae vel incisae; folia membranacea usque medium 3-lobata basi subcordata lobis oblongo-lanceolatis argute acuminatis serratis supra pilosula deinde glabrata, petiolo 3-4 glandulis incon- spicuis praedito; bracteae liberae ovato-lanceolatae serrulatae; calycis tubus cylindricus glaber, sepalis petalisque ovato-oblongis obtusis tubo paulo brevioribus; corona circulum minutorum tuberculorum reducta; operculum deflexum recurvatum; ovarium brunneosericeum.

Herbaceous vine, densely brownish or grayish tomentose throughout except the upper side of the leaves and the flowers; stem terete; stipules broadly ovate-reniform, about 1 cm. long and 1 cm. wide at the base, scarcely oblique, closely semiamplexicaul, irregularly dentate or laciniate at the apex, the divisions aristate; petioles 2-2.5 cm. long, bearing 3 or 4 obscure glands; leaves 6-9 cm. long, 8-9 cm. wide, 3-lobed about to the middle (lobes oblong-lanceolate, 2.5-3.5 cm. wide, sharply acuminate, the lateral somewhat ascending), conspicuously serrate, subcordate, membranous, the upper surface sparingly pilosulous glabrate; peduncles 12-15 cm. long, rather stout; bracts free to base, ovate-lanceolate, about 3 cm. long, and 1.7 cm. wide, acuminate, serrulate, sessile, borne at the apex of the peduncle; calyx tube cylindric, 7-7.5 cm. long, 1-1.3 cm. wide when pressed flat, slightly enlarged at baso, otherwise of uniform diameter, glabrous; sepals ovate-oblong, 6-6.5 cm. long, about 2.5 cm. wide, obtuse, dorsally awned just below the apex (awn very slender, about 5 mm. long), deep salmon-rose, paler within, the midnerve with a pale green center; petals similar and subequal to the sepals; corona a single series of minute tubercles ; operculum borne near the base of the tube, about 3 mm. long, dependent, the margin recurved; ovary ovoid, densely light-brown-sericeous.

Type in the Herbarium of the Chicago Natural History Museum, No. 1204733, collected in the vicinity of Achupallas, Province of Santiago-Zamora, Ecuador, at an altitude of 2,500 m. in a forest along a stream on October 8, 1943, by Julian A. Steyermark (No. 54493). Duplicate at US.

In the monograph the new species keys out to P. jamesonii because of the large stipules; however, these are far more deeply fringed in P. jamesonii. Moreover, P. jamesonii is glabrous throughout and has shorter peduncles and deeply fringed bracts. In leaf shape, indument, and length of the peduncles, the new species suggests P. ampullacea.


162. Passiflora matthewsii (Mast.) Killip

Masters placed this species among the tacsonias with united bracts, lobed leaves, and narrowly linear stipules, though in the description of the species, which was given at some length in the key, he did not specifically mention the stipules. A specimen recently collected at the type locality, Chachapoyas, Peru, by Metcalf (Goodspeed Expedi- tion No. 30784) has foliaceous, semiovate stipules which are deeply pinnatisect into filiform segments, the body of the stipules sometimes being reduced to a mere band surrounded by the filiform segments. Otherwise the specimen agrees perfectly with a photograph that I took of the type at the Kew Herbarium and with the amplified descrip- tion given in my monograph. I believe that both Masters and I mistook either the filiform segments or abortive tendrils for stipules, that the stipules of P. matthewsii had mostly fallen off the type material at Kew and the British Museum, and that therefore P. matthewsii should be placed next to P. roseorum (species 164).

A. H. G. Alston has kindly examined for me the Matthews specimen at the British Museum, and writes: "The stipules appear to have fallen off our specimen, but there are sometimes young tendrils which could easily have been taken for stipules. ... On the young parts there are, however, some filiform hairy segments, which could easily be the segments of a laciniate stipule."

166a. Passiflora loxensis Killip & Cuatr., sp. nov. Plate 7

Suflrutex scandens, caule angulato glabro; stipulae coriaceae ovatae fimbriato-dentatae; folia coriacea glabra trilobata lobis triangulari- ovatis acutis serratis, subtus reticulo venorum valde elevato con- gesteque incrassato; bracteae ad medium connatae; calycis tubus cylindricus glaber, sepalis petalisque oblongis tubum subaequilongis; corona tuborculata; operculum deflexum paulo laciniatum recurva- tum; ovarium glabrum.

Vine; stem aeute-angular, glabrous, lustrous; stipules coriaceous, glabrous, ovate or lance-ovate, acuminate, oblique and cordate at base, amplexicaul, fimbriate-dentate, 10-20 mm. long, 4-8 mm. broad; petioles 1-2 cm. long, bearing 3-4 sessile or short-stipitate glands about 1 mm. thick; leaf-blades 4-6 cm. long, 5.5-9 cm. broad, deeply 3-lobed (lobes triangular-ovate, acute, 2-3.5 cm. long, 1.5-2.8 cm. wide, the lateral ones subascending), truncate or subrounded at base, callous-serrate, coriaceous, glabrous, above green, rugulose, the nerves impressed, the minute reticulate veins prominulous, beneath paler, the 3 primary nerves thick and prominent, the secondary nerves prominent, the veins thick forming an incrassate reticulum separating very narrow, deep, irregular alveolae; peduncles 2.5-4.5 cm. long, subangular, stout; bracts subcoriaceous, 5-6 cm. long, connate to


about the middle (the free part oblong, abruptly acuminate, entire, 1.4-2 cm. wide), outside glabrous, slightly nervate and reticulate, inside more or less lanuginous or pilose and conspicuously venose; flowers lilac-blue; calyx tube cylindric, 7-8 cm. long, about 1 cm. broad when pressed, of uniform diameter, glabrous; sepals oblong, obtuse, aristulate below the apex, 6.5-8 cm. long, 1.8-2.6 cm. wide; petals subequal to the sepals; corona tuberculate; operculum depend- ent, about 7 mm. long, the margin recurved, more or less fimbriate; gynophore finely spreading-pubescent ; ovary fusiform, glabrous; immature fruit obovoid-oblong, about 6.5 cm. long, 2 cm. broad; seeds oblong-ovate, 7-8X4-5 mm., alveolate.

Type in the U.S. National Herbarium, No. 1909202, collected at Horta-Naque, Province of Loja, Ecuador, at an altitude of 3,200 m., on Nov. 8, 1946, by Reinaldo Espinosa (No. E965). Supplementary sheet, US No. 1909586.

P. loxensis belongs to the subgenus Tacsonia and is closely related to P. cumbalensis (Karat.) Harms. It differs in the laciniate-dentatc, coriaceous stipules, in the more coriaceous leaves and bracts, and especially in the structure of the lower surface of the leaves. The thick veins on the lower surface form a very close reticulum that leaves very thin but deep and irregular pits between the turgid network.

170. Passiflora psilantha (Sodiro) Killip

This Ecuadorean species has been found in cultivation near Sons6n, Antioquia, Colombia, by Gutierrez & Barldey (No. 18C294). The specimen is deposited in the Herbarium of the Facultad de Agrononria, Medellm, Colombia.

173. Passiflora mixta L.f.

On page 297 of my monograph, variations of pubescence in this wide-spreading species are noted thus:

(1) Leaves, calyx tube, and ovary glabrous.

(2) Leaves glabrous, calyx tube and ovary pubescent.

(3) Under surface of leaves, calyx tube, and ovary pubescent. No formal varietal names were assigned to these variants though a fourth, P. mixta var. eriantha, was treated as a variety.

Two specimens received subsequent to the publication of my monograph exhibit other combinations in the occurrence of indument, and are worth noting:

(4) Under surface of leaves pubescent, calyx tube glabrous, ovary pubescent: Ecuador: Canar: Tipococha, altitude 3,200 m., Penland cfc Summers 1006 (US).

(5) Leaves and calyx tube glabrous, ovary pubescent: Bolivia: La Paz: Charazani, Province of Muiiecas, altitude 2,700 m., Cdrdenas 3886 (US).


177. Passiflora antioquiensis Karst.

In the description given of this species in my monograph, notes on the peduncle were unfortunately omitted. It is worthwhile also to amend the description in certain details in view of recent collections thus:

Leaves unlobed, up to 18 cm. long and 8.5 cm. wide, asymmetrically 2-lobed or 3-lobed to within 2 cm. of the base; peduncles very slender, almost filiform, 17-60 cm. long, pendulous; flowers up to 14 cm. wide when expanded.

Additional specimens examined are:

Colombia: Antioquia: Yarumal, Daniel 1916 (US). Cauca: El Tambo, alt. 2,500 m., von Sneidern 1442 (S.XJS).

180a. Passiflora macropoda Killip sp. nov. Plate 8

Caulis scandens robustus ruf o-subvelutinus ; stipulae cordato-ovatae acutae inciso-fimbriatae glabrae; folia subcoriacea pilosula ad medium trilobata basi subcordata lobis ovatis acutis serratis ; bracteae liberae ovato-lanceolatae subcordatae inciso-fimbriatae; calycis tubus tubu- loso-campanulatus, sepalis anguste lanceolatis pilosulis apice longe mucronatis quam tubo duplo longioribus, petalis leviter brevioribus; corona 3 seriebus filamentis filiformibus et interiore brevi membrana horizontali subcrenata; operculum deflexum recurvum; gynophorum tener; ovarium velutinum.

Vine; stem terete, stout, about 5 mm. in diameter, rufo-velutinous ; stipules cordate-ovate, 2-3 cm. long, 1.5-2 cm. wide, acute, strongly oblique at the base, Iaciniate-fimbriate, membranous, glabrous; petioles stout, 1.5-3 cm. long, bearing 2 or 3 short stipitate glands at the apex and 1 or 2 just below the apex, these partially concealed in a dense tomentum; leaf -blades 3-lobed about to middle, 10-12 cm. along the midnerve and the lateral nerves, 16-18 cm. between the apices of the lateral lobes (lobes about 5 cm. wide, aristate), cordulate at base, serrulate, subcoriaceous, minutely pilosulous on the nerves above and on the principal nerves beneath cano-puberulent or rufo- puberulent on the secondary nerves and the veins beneath ; peduncles solitary, stout, 12-14 cm. long, densely rufo-velutinous; bracts free to base, ovate-lanceolate, 4-5 cm. long, about 2 cm. wide, cordulate, laciniate-fimbriate, the teeth toward the base the largest; calyx tube slenderly campanulate, about 2 cm. long, and 1.3 cm. in diameter; sepals narrowly lanceolate, 4 cm. long, 1 cm. wide, white within, dorsally keeled toward the apex, the keel terminating in a filiform awn about 8 mm. long; petals narrowly lanceolate, 3.5-4 cm. long, white; corona in 4 series, the outer 3 ranks irregular, filamentose, the filaments filiform, about 1.5 cm. long, close to these ranks a short, thick, horizontal, subcrenulate membrane; operculum dependent,


about 8 mm. long, the margin recurved, short filamentose; gynophore slender, about 3.5 cm. long; ovary ovoid, white- velutinous.

Type in the U.S. National Herbarium, No. 1803911, collected near Unduavi, Department of La Paz, Province of Nor Yungas, Bolivia, at an altitude of 3,000 m., in forest along roadside, on Apr. 15, 1939, by W. J. Eyerdam (Goodspeed Expedition No. 25379). Duplicates at University of California and Chicago Natural History Museum.

This species belongs to the subgenus Granadillastrum and because of its laciniate-fimbriate bracts and general flower structure is evidently nearest P. weberbaueri and P. trisecta. Both of these have trifoliate leaves. The indument and general shape of the foliage are suggestive of P. peduncularis.

189. Passiflora racemosa Brot.

In describing the bracts of this species I said in the monograph: "bracts setaceous (?), soon deciduous." This statement is in error. In Chase 10125, which was cited, bracts are present, and Masters' plate in the Flora Brasiliensis shows a bract. They are ovate, about 1 cm. long and 8 mm. wide, tapering slightly at the base, thin-mem- branous, and pink, and are borne close to the base of the flower.

P. G. Greenway has recently sent me for study a superb lot of passionflowers from the Herbarium of the East African Agricultural Research Station, Amani, Tanganyika, nearly all of which are from plants grown at the Station and are of New World origin. Among these, No. 5942, collected by Greenway, is P. racemosa. The flowers are borne singly in the axils of fully developed leaves and are not in a raceme or on long branches with much reduced leaves. This plant may be of hybrid origin, or perhaps in a natural state the flowers may sometimes be solitary.

200. Passiflora seemannii Griseb.

This species has a rather unusual range of distribution. At the time that my monograph was prepared, the species was known in its native state by several collections from Panama and from scattered localities in Colombia. The following additional collections have sub- sequently been received at the U.S. National Herbarium:

Colombia: Cundinamarca: San Francisco, alt. 1,550-1,580 m., Garcla- Barriga 11051. El Meta: Villavicencio, alt. about 500 m., Killip 34445, 35692; Cuatrecasas 4673. Venezuela: Amazonas: Yavita, L. Williams 14000.

209. Passiflora capparidifolia Killip

Material from British Guiana and the Rio Branco of Brazil was cited as this species in my monograph. The species is now represented in the U.S. National Herbarium by the following Venezuelan collec- tions :


Bolivar: Between Rio Torono and Salto de Auraima, edge of forest along Rio Paragua, alt. 275 m., Killip 37539. Selvas de galeria del Rio Uari, Gran Sabana, Tamayo 3154.

The first of these collections was in fine flower and the second one bore young fruit. The description in the monograph may be amplified thus:

Bracts distinct to the base, broadly ovate, up to 5 cm. long and 3 cm. wide; sepals greenish white without, red-brown within; petals deep pinkish red; outermost corona filaments about 1.2 cm. long, banded with blue and white in upper part, red and white in lower part, the second series three times longer, blue and white in the upper two- thirds, red and white in the lower third; young fruit ovoid, about twice as long as wide, finely pilose, obcordate, about 5 mm. long and 3.5 mm. wide, reticulate.

210. Passiflora popenovii Killip

At the time that my monograph was prepared the only specimens of this species available to me were from Ecuador. The species was also collected in 1938 at El Tambo, Department of Cauca, Colombia, alt. 1,700 m. by K. von Sneidern (No. 1444), and was deposited in the Riksmuseet, Stockholm, and in the U.S. National Herbarium.

211a. Passiflora brevifila Killip, sp. nov. Plate 9

Frutex scandens omnino glaber, caule subquinqueangulato; stipulae anguste lineares subfalcatae; folia subcoriacea integra lanceolata vel oblongo-lanceolata acuminata basi rotundata; bractcae liberae lanceolatae minute cuspidatae integrae basi subtruncatae, petiolo prope apicem 4-6 glanduloso; calycis tubus campanulatus, sepalis petalisque similibus oblongis obtusis; corona filamentis uniseriatis liguliformibus ad apicem leviter dilatatis; operculum fimbriatum fila- mentis rigidulis ad basim paulo incrassatis.

Woody vine, glabrous throughout; stem subquinquangular; stipules narrowly linear, 5-6 mm. long, about 0.8 mm. wide, subfalcate, coriaceous; petioles 1-2 cm. long, bearing 4-6 minute glands near the apex, the glands short-stipitate or nearly sessile; leaves lanceolate or oblong-lanceolate, 12-14 cm. long, 5-5.5 cm. wide, acuminate, rounded at base, entire, penninerved (principal lateral nerves 6 or 7 to a side, arcuate-ascending), conspicuously reticulate beneath, sub- coriaceous, concolorous, lustrous beneath; peduncles solitary, 2.5-3 cm. long, bracts free to base, lanceolate, 5-7 cm. long, 2-3 cm. wide, acuminate, minutely cuspidate, sub truncate at base, entire, similar in texture to the leaves; flowers about 6.5 cm. wide; calyx tube campanu- late; sepals and petals similar, oblong, about 2.5 cm. long and 9 mm. wide, obtuse, apparently white; corona filaments in a single series, narrowly liguliform, slightly thickened at apex, 1 cm. long; operculum


a fringe of rather stiff filaments about 3 mm. long, slightly thickened at base; limen a thick ridge close to the operculum; floor of calyx tube smooth; ovary ovoid, green; fruit globose, about 6 cm. in diameter, the exocarp brittle, orange; seeds obovate, strongly flattened, 9-10 mm. long, 5 mm. wide, coarsely reticulate, tridentate at the apex, the middle tooth the more prominent.

Type in the U.S. National Herbarium, No. 1643741, collected at Vara Blanca de Sarapiquf, north slope of Central Cordillera, between Poas and Barba Volcanoes, Costa Rica, at an altitude of 1,890 m., in February 1938, by A. F. Skutch (No. 3591).

This species belongs to the subgenus Oranadilla series Laurifoliae, where it is the only one with a single-ranked corona. The other representative of the series in Central America, P. ambigua, not only has a 5-ranked corona but the ovary is densely brown-tomentose. Doubtless the closest relative is P. nigradenia, of Bolivia, in which the corona is 2-ranked and the ovary finely sericeous.

213a. Passiflora killipiana Cuatr., sp. nov. Plate 10

Planta breviter hirtella vel pubescens, pilis patulis ruf escentibus ; stipulae lineari-subulatae; folia tenuiter subcoriacea ovato-oblonga vel elliptico-oblonga basi subrotundata apice acute acuminata margine remote minuteque crenulata, penninervia, nervis utrinque ferrugineo- hirtellis vel tomentulosis, venulis reticulatis utrinque prominulis, petiolo brevi subapicem biglanduloso; bracteae liberae obovato- oblongae apice rotundatae sursum profunde dentatae; calycis tubus campanulatus, sepalis oblongis subapicem corniculatis, petalis brevioribus; corona filamentis 6-7 seriatis exterioribus liguliformibus valde longis, interioribus mediocribus, intermediis brevibus; operculum membranaceum profunde fimbriatum; ovarium velutino-tomentosum. Plant hirtellous or pubescent nearly throughout, the hairs reddish, short, spreading; stem terete, slightly striate; stipules linear-subulate, 10-13 mm. long, 1 mm. wide, ferruginous-hirtellous, soon deciduous; petioles 7-11 mm.