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Manual de Plantas de Costa Rica

Main | Family List (MO) | Family List (INBio) | Cutting Edge
Draft Treatments | Guidelines | Checklist | Citing | Editors

Draft Treatments

CACTACEAE
By B. E. Hammel
English, work in progress: placed 1/Apr./2000

1 Plants terrestrial or sometimes clambering on other vegetation, but rooting only at the base (or at nodes); stems with spines (except a few cultivated Opuntia).

2 Leaves and stems normal (leaves broad and thin, stems cylindrical and unribbed)…Pereskia

2' Leaves and stems cactoid (leaves small points seen only on new growth, or lacking, stems variously ribbed or broadly flattened).

3 Stems (or side branches from them) broadly flattened, the branch segments more or less obovate; fine hairs at base of spines very brittle and barbed…Opuntia

4 Plants spineless; cultivated.

5 Largest pads mostly < 25 cm long, <15 cm wide; fls red...O. cochenillifera

5' Largest pads > 30 cm long, >20 cm wide; fls. yellow…O. ficus-indica

4' Plants with spines; native or cultivated.

6 Spines yellowish, both hair-spines and spines present; very rare, native (Palo Verde); fls. red?…O. lutea

6' Spines white to gray, only spines present; relatively common, native or cultivated; fls. yellow.

7 Plants with cylidrical central trunk 2 m or more tall, with numerous cylindrical lateral branches from which the pads arise; cult.…O. brasiliensis

7' Plants without trunk or, if trunk present, < 2 m, the lateral branches all formed by the connected pads.

8 Largest pads > 30 cm long, >20 cm wide; plants erect; cult. O. ficus-indica

8' Largest pads < 20 cm long, <10 cm wide; plants often prostrate; native...O. guatemalensis

3' All stems basically cylindrical, but variously ribbed or winged; fine hairs at base of spines, if present, not barbed.

9 Plants 2 m or more tall, erect.

10 Stems 25-35 cm wide, 10+ ribbed, erect; stem tips with horizontal, zig-zag, glaucous lines…Stenocereus aragonii

10' Stems ca. 15 cm wide, 4--6 winged, often arched and clabering on other vegetation; stem tips unadorned…Acanthocereus tetragonus

9' Plants < 50 cm tall, decumbent or erect.

11 Spines recurved; stems more or less globose, unbrached, erect; fls. clustered at very tip of stem, among wooly hairs…Melocactus curvispinus

11' Spines erect; stems elongate, cylindrical and prostrate; fls. growing along stem…Peniocereus hirschtianus

1' Plants epiphytic or (occasionally) clambering on rocks, usually producing roots along the stem, as well as at base; stems with or without spines.

12 Stems regularly di- to poly-chotomously branched; seeds narrowly ovate; fls. without tube, the limb < 5 mm wide, white; fr. globose, ca. 5 mm diam, lacking bracteoles, translucent white…Rhipsalis

13 Stems cylindrical; Atl. lowlands and wet forest of (S) Pac. lowlands…R. baccifera

13' Stems 4-angled; moist forest, meseta central…R. micrantha

12' Stems irregularly and not di- to poly-chotomously branched, broadly flattened or 3 or more ribed or winged; seeds +- pear-shaped; fls. with tube ca. 0.5--30 cm, rarely nearly lacking, the limb pink, red, or white; frs. usually ovoid and much larger, with bracteoles, usually red or green, not white.

14 Ovary bracts without stiff spines; stems either broadly 3-angled, or flattened.

15 Ovary bracts large (mostly > 0.5 cm), overlapping, without wool or hair-spines; fls. broadly funnel-shaped; stems thinly to thickly 3-winged, usually with 2--6, spines in the aeroles…Hylocereus

16 Stem wings thick (3 mm or more in fresh material), the stems usually more or less glaucous; fls. ca. 30 cm; plants apparently native only on Pac. Slope.

17 Bracteoles of ovary 0.5--3 cm, imbricate; spines 2--4 mm, 3--6/areole, mostly black; stigma lobes not cleft; known only from N half of country, coastal and inland…H. costaricensis

17' Bracteoles of ovary 0.5--1 cm, distant; spines 1--2 mm, usually 1 or 2/areole, reddish brown; stigma lobes apically cleft; known only from S half of country, inland…H. monacanthus

16' Stem wings thin (ca. 2 mm at most), green, not glaucous; stout spines often lacking; plants apparently native only on the Atlantic slope.

18 Wings of stems with prominent rounded projections 1--2.5+ X 1.5--2 cm; fls. huge (30--37 cm), the inner tepals white, the upper bracteoles papery thin and becoming chartaceous; stout spines almost always lacking…H. calcaratus

18' Wings of stems lacking prominent projections, more or less regularly undulate; fls. relatively small (8--)9--10 cm, the inner tepals red or pink, the upper bracteoles more or less thickly succulent; 1--3 stout spines present or not…H. stenopterus

15' Ovary bracts small (mostly < 0.3 cm); fls. trumpet-shapped or with essentially no tube; stems broadly flattened, the aeroles without stiff spines.

19 Fls. at least 10 cm; fls. trumpet-shaped…Epiphyllum

20 Areoles with abundant felt, the lower ones of stem often with several, weak hair-spines; stems drying green, the margins +- deeply crenate and often abruptly rounded to the sinus apically; tepals ca. 6--8 cm…E. lepidocarpum

20' Areoles lacking felt.

21 Stems (phylloclades) 8--25 cm wide, usually drying black or gray, the margins shallowly to deeply lobed, but +- regularly; tepals 9--14 cm.

22 Areoles of stem sinuses with conspicuous, protruding flaps, ca. 2 mm; stems, thick and stiff (live material), very deeply to shallowly lobed, the margines distinctly convex; areoles lacking hair-spines; fl. buds +- in same plane as stem, but pointing away from its edge…E. grandilobum

22' Areoles of stem sinuses without conspicuous flaps; stems succulent but rubbery flexible, shallowly lobed, the margins usually nearly flat; areoles (especially of ovary) often with hair-spines; fl. buds +- pressed against the lower surface of stem…E. thomasianum

21' Stems 3--6.5 cm wide, mostly drying yellowish green or gray, the +- shallow lobes slightly broader apically; tepals ca. (3.5--)4--6 (--7)cm.

23 Reticulate venation of stem lamina clearly visible in dry material, the surface often drying shiny; seed surface smooth…E. cartagense

23' Reticulate venation of stem lamina not visible in dry material, the surface usually drying dull; seed surface conspicuously pitted…E. phyllanthus

19' Fls. < 4 cm +- funnel-shaped, without tube, or trumpet-shaped…Disocactus

24 Fls. with no apparent tube, funnel-shaped or rotate, the tepals ca. 6 mm, the outer ones translucent white to pink; flattened portion of stems relatively thin, drying +- green.

25 Reticulate venation of stem lamina clearly visible in dry material, the surface drying shiny; stems to 7 cm wide; fls. funnel-shaped…D. acuminatus

25' Reticulate venation of stem not visible in dry material, the surface drying dull; stems mostly < 3 cm wide; fls. rotate…D. ramulosus

24' Fls. with an obvious tube, narrowly funnelform or trumpet-shaped, outer tepals reddish cream or lavender, the tepals ca. 8--15 mm; flattened portion of stem relatively thick, usually drying dark gray or tan.

26 Fls. lavender; frs. ovoid; seeds dull, minutely pitted; known only from moist forest, Guanacaste, very rare (in CR)…D. amazonicus

26' Fls. pale reddish cream; frs. globose to ovoid; seeds smooth, shiny; known only from wet Pacific forest (from Carara south), common…D. himantocladus

14' Ovary bearing +- stiff spines or at least tuberculate and with hair-spines (check, eg., Weberocereus biolleyi); stems various, often with spines

27 Fl. tube ca. 10 cm long (fls. trumpet-shaped); stems cylindrical (5--12 ribbed or winged)…Selenicereus

28 Stems spiney, ca. 5--15 (or more?) cm diam., including the 5-7 broad wings, tightly clinging to substrate…S. testudo

28' Stems without spines, ca. 1.5 cm diam, with numerous low ribs, loosely climbing and pendulous…S. wercklei

27' Fl. tube mostly much less than 10 cm (fls. +- funnel-shaped); stems all of the above, often with hairs, bristles or spines in the areoles; seeds minutely pitted (except W. tonduzii)…Weberocereus & "Nopaloxchia horichii??"

29 Stems mostly 10--15 cm wide, flattened.

30 Lobes of stem very deep; frs. with spines, the pulp white…W. imitans

30' Lobes of stem very shallow; frs. lacking spines, the pulp reddish…W. bradei

29' Stems < 6 cm wide, cylindrical or variously ribbed to flattened or 3 or 4 winged.

31 Stems (2)3--4(--5) cm wide, broadly flattened or 3-winged.

32 Inner tepals reddish pink, the fls. ca. 8 cm (--16!!!?--not true on the plants Horich gave me, ca. _ that size. This measurement from the type description), the tube with a few stiff bristles basally; stems mostly thickly flattened to 3-angled, the areoles with conspicuous wool and several stiff bristles; fr. flesh pinkish white…"Nopaloxchia horichii"

32' Inner tepals cream, the fls. ca. 6--8 cm, the tube mores or less covered with short spines; stems mostly thinly 3 winged, the areoles with or without spines; fr. flesh yellow, fragrant and sweetly acidic; seeds smooth; …W. tonduzii

31' Stems < 2 cm wide, +- cylindrical, smooth to variously ribbed or angled; fr. flesh dark magenta, tasteless and odorless; fls. ca 5--7 cm.

33 Areoles of stems with numerous yellowish hair-spines as well as spines…W. trichophorus

33' Areoles of stems without hairs or the hairs white.

34 Spines abundant on stems, hairs lacking; fls. 6--7 cm…W. tunilla

34' Spines and hairs often lacking on stems; fls 5--6 cm.

35 Stems mostly smooth…W. biolleyi

35' Stems mostly sharply 3-4 angled…W. panamensis

Acanthocereus

Acanthocereus tetragonus (L.) Hummelinck.

Terrestrial, long-stemmed, up to 4 m, often forming clumps in coastal thickets, the stems arching and sometimes more or less recumbent on other vegetation; stems mostly 3 or 4 winged, the wings 3--5 cm wide (i.e. from base to marging), the margins shallowly crenate-lobed, 4--6 cm between areoles, the areoles with dense wool and 10+ stout spines 0.2--7 cm, lacking bracts. Fls. trumpet-shaped, 13--22 cm, tepals 3--5 cm, the external ones green (sometimes tinged violet), inner ones white; ovary with numerous short (ca. 2mm) bracteoles subtending dense wool and few short (1--3 mm) spines), the tube also with few such bratcteoles. Fr. gobose, spiney, ?? cm, reddish-purple, the flesh ??; seeds ?shaped, ?? mm, the surface?, black.

Dry, coastal thickets and canyons, 0--800 m; Pacific slope, Guanacaste to Valle Central and disjunct to the Rio Terraba Canyon. (Hammel 18494, INB)

This is one of our most commonly cultivated native cacti. The usually long spines and few, thin stem wings on this ererct to arching terrestrial species, are distinctive. The disjunct population along the lower reaches of the Terraba river appears to have generally shorter spines (to 2 cm) and fls. (to13 cm) than the more northern populations.

Disocactus (sensu Kimnach)

Mostly pendulous, spineless epiphytes with flat stems and with small (<3 cm? long) fls. but with clearly visible tube; frs. globose or broadly ovoid.

Disocactus acuminatus (Cufod.) Kimnach, Cact. Succ. J. (Los Angeles) 33: 14. 1961; Pseudorhipsalis acuminata Cufod., Arch. Bot. Sist. 9(3-4): 196. 1933; Rhipsalis acuminata (Cufod.) Standl. Disocactus horichii Kimnach.

Cladodes 10--50 x 2--7 cm, the apex mostly narrowly acuminate, rarely rounded, the lamina drying +- translucent, often shiny, green or grayish green; margins shallowly serrate to crenate, the lobes (areole to areole x bottom of sinus to apex of lobe, measured at wides part of cladode) 10--30 x 1--2 (--5) mm, broadest at their middle or towards their apex, usually convex. Bracts of areoles ca. 0.2? mm, the areoles essentially naked (on older stems). Fls. rotate or funnelform, ca. 1--1.6 cm (incl. ovary); tepals 6--1.2 cm, white the outer ones sometimes tinged pink; ovary with a few bracteoles (0.2--0.3 mm) subtending minute hairs (<0.1 mm). Fr. globose, ca. 5--6(--8) mm, red or pink; seeds pear-shaped, 1.6--1.8 mm, the surface smooth (slightly pitted near neck) and dull, reddish brown to black.

Wet forest, 40--900 (-1400) m; Cordillera Guanacaste to Cordillera Volcanica Central, and Llanuras de San Carlos on the Atlantic slope, Fila Bustamante on the Pacific slope..

(incl. material from La Selva--Hammel 8680 & D. Smith 58). Thin leaves drying with very obvious reticulate venation. Fresh leaves described as "somewhat rubbery, not succulent…brittle when bent…." Horich s.n., given to me by Horich as D. horichii and claimed by him to be from the type plant, flowered (and was vouchered) in the MO greenhouse on 23 May 1994, is D. acuminatus.

Disocactus amazonicus (K. Schum.) D. R. Hunt, Cact. Succ. J. Gr. Brit. 44(1): 2. 1982; Wittia amazonica K. Schum., Monatsschr. Kakteenk. 13: 117. 1903.

Cladodes 20--30+ x 2--6+ cm, the apex +- rounded to acuminate, the lamina drying dull and dark gray; margins shallowly crenate, the lobes 15--30 x 1--2 mm, broadest towards their apex. Bracts of areoles ca. 0.2 mm, the areoles with very short (0.2 mm) wool. Fls. +- tubular, 2.5--3(--4) cm; tepals ca. 1 cm, lavender; ovary with a few bracteoles (ca. 0.5 mm) subtending minute hairs. Fr. +- ovoid, 1--1.5 cm, pink to red; seeds pear-shaped, 1.6--1.8 mm, surface ruggose and sometimes slightly pitted, dull black.

Nic.--Peru.

Guanacaste, Carmona, Cerro Azul, 900 m, fl. Feb 28 '86 "Flores de color rosado con lila muy vistosas." Very broad leaves, much like an Epiphyllum, magenta more or less tubular fls. (L. D. Gómez et al 23346, CR; Horich s.n., Cerro el Pando de Nicoya)

Disocactus himantocladus (Rol.-Goss.) Kimnach, Cact. Succ. J. (Los Angeles) 33: 14. 1961; Rhipsalis himantoclada Rol.-Goss., Bull. Soc. Bot. France 55: 694. 1908. Disocactus lankesteri Kimnach.

Cladodes 20--45(--60) x (1.5--)3--4.5 cm, the apex rounded to acuminate, the lamina drying dull and dark gray to tan; margins shallowly crenate, the lobes mostly 15--20 x 1--2 mm, broadest towards their apex or at their middle, convex to straight. Bracts of areoles 3--5 mm, the areoles (of mature stems) essentially naked. Fls. trumpet-shaped, 1.5--2.5(--3.3) cm; tepals ca. 0.8--1(--1.5) cm, the outer yellowish pink, the inner white; ovary with a few bracteoles (ca. 0.5 mm), hairs lacking. Fr. +- globose, ca. 1 cm, reddish; seeds broadly pear-shaped, 1.3--1.6 mm, surface minutely pitted, dull, dark reddish brown to black.

Wet but somewhat seasonal forest, 0--650 m; Pacific slope from Valle de Parrita to the Osa Peninsula, also (rare) in the Valle de El General. CR & Pan.

Much material from CR was formerly misidentified as D. biformis, which does not occur here. Broad stems drying dark and dull; pinkish cream fls. with a rather long tube (ca. 1 cm). Fresh leaves stiff and brittle. Disocactus lankesteri, isotype in fl. at MO, has the same unusual pollen in tetrads of D. himantocladus, and is in all other ways indistinguishable from said species.

Disocactus ramulosus (Salm-Dyck) Kimnach, Cact. Succ. J. (Los Angeles) 33(1): 14. 1961; Cereus ramulosus Salm-Dyck, Hort. Dyck. 340. 1834; Rhipsalis ramulosa (Salm-Dyck) Pfeiff. Rhipsalis coriacea Polak.

Cladodes 10--27 x 1--2.5 cm, the apex rounded to acuminate, the lamina drying dull and greenish tan; margins shallowly crenate, the lobes mostly 15--20 x 1--2 mm, broadest towards their apex. Bracts of areoles ca. 0.5 mm, the areoles naked. Fls. rotate, ca. 1 cm; tepals 0.5--0.6 cm, white; ovary with a few bracteoles (ca. 0.5 mm), hairs lacking (except rarely scant hairs at base of fl). Fr. globose, 0.5--0.6 cm, white; seeds pear-shaped, 1.2--1.3 mm, surface smooth, dull, black.

Wet forest, 700-1350 m; Cord. Tilaran (both slopes) to Rio Reventazon.

Narrow stems, small white fls. with basically no tube.

Epiphyllum

Flat- and broad-stemmed, spineless, pendulous epiphytes with large fls. (mostly> 7 cm?); frs. narrowly ovoid.

Epiphyllum cartagense (F. A. C. Weber) Britton & Rose, Contr. U.S. Natl. Herb. 16: 256. 1913; Phyllocactus cartagensis F.A.C. Weber, Bull. Mus. Hist. Nat. (Paris) 8: 462. 1902.

Cladodes 13--45 x (1.5--)2.5--6.5 cm, the apex mostly rounded to acute, rarely acuminate, the lamina often drying dark, greenish tan and somewhat shiny, the reticulate venation usually obvious; margins markedly crenate, the lobes mostly 25--55 x 4--15 mm, broadest towards their apex. Bracts of areoles ca. 0.2--0.5 mm, the areoles with very short hairs. Fls. trumpet-shaped, 16--19(--21) cm; tepals 5--6.5 cm, the outer pinkish, the inner white; ovary with a few triangular bracteoles (0.5--2 mm), subtending scant, short hairs. Fr. narrowly ovoid, (4--)5--8 cm, obscurely 4or 5 angled, red with white pulp; seeds pear-shaped, 3--4 mm, surface faintly rugose, dull to somewhat shiny, black.

Wet forest, 700--1700 m; both slopes from Cordillera Guanacaste to Cordillera Talamanca. Endemic.

The obvious reticulate venation and often somewhat shiny surface of the cladodes (dry material) help distinguish this species from other flat-stemmed cacti in CR.

Epiphyllum grandilobum (F. A. C. Weber) Britton & Rose, Contr. U.S. Natl. Herb. 16: 257. 1913; Phyllocactus grandilobus Weber, Bull. Mus. Hist. Nat. Paris 8: 464. 1902. Epiphyllum gigas Woodson & Cutak

Cladodes 40--70+ x 8--12(--25) cm, the apex rounded to broadly acute, the lamina drying dull, usually light greenish or grayish tan; margins shallowly crenate (to very deeply lobed), the lobes 40--70 x (2--)4--20(--60) mm, mostly broadest towards their middle. Bracts of areoles 1--2(--4) mm, the areoles naked. Fls. funnel- or trumpet-shaped, 28--38 cm; tepals 10--15 cm, the outer greenish yellow the inner cream to white; ovary with a few bracteoles (2--15 mm) subtending scant wool. Fr. ovoid, ca. 5 cm; seeds pear-shaped, 2.5--3 mm, surface slightly pitted, dull, black.

Wet forest, 20--1100 m; Cordillera Tilaran, Llanura de Tortuguero, Osa Peninsula. Nic.--Pan.

The very conspicuous areolar bacts (in the sinus' of stem lobes), help distinguish this species, vegetatively from all other flat-stemmed cacti in CR. In life, the cladodes are the thickest of CR Epiphyllum, but not as thick as those of Weberocereus bradei. The one fr. collection has the fr. attached directly to the woody cylindrical stem, not to the edge of a cladode.

Epiphyllum lepidocarpum (F. A. C. Weber) Britton & Rose, Contr. U.S. Natl. Herb. 16: 257. 1913; Phyllocactus lepidocarpus F.A.C. Weber, Bull. Mus. Hist. Nat. (Paris) 8: 462. 1902.

Cladodes 30--75 x 3--5.5 cm, the apex rounded to acute, the lamina drying dull and greenish tan; margins moderately crenate, the lobes mostly 25--40 x 5--7 mm, broadest towards their apex. Bracts of areoles ca. 0.5 mm, the areoles usually distinctly wooly, often with a few hair-spines. Fls. funnel-form to trumpet-shaped, 14--17(--20) cm; tepals 5.5--7(--8) cm, the outer rose, the inner pale rose to creamy white; ovary with a few bracteoles (ca. 2--3 mm), subtending wool and occasional hair-spines. Fr. ovoid to globose, 4.5--5 cm, purplish pink; seeds pear-shaped, 1.5--2 mm, surface smooth, shiny, reddish black.

Wet forest, 700--2050 m; Cordillera de Tilaran to Cordillera Talamanca and Cerros Escazu. CR & Pan.

The areoles usually have obvious wool and sometimes hair-spines and the apparently quite succulent cladodes dry greenish tan. The epidermis of the cylidrical portions of stems often dries yellowish tan and brittle.

Epiphyllum phyllanthus (L.) Haw., Syn. Pl. Succ. 197. 1812; Cactus phyllanthus L. Sp. Pl. 469. 1753. Epiphyllum pittieri (F. A. C. Weber) Britton & Rose.

Cladodes 17--100+ x 3--7 cm, the apex rounded to acute, the lamina drying dull and grayish to greenish tan; margins moderately crenate, the lobes mostly 25--50 x 3--8 mm, broadest towards their apex, rounded to often concave. Bracts of areoles ca. 0.2 mm, the areoles with very short wool. Fls. trumpet-shaped, 10--14.5(--18) cm; tepals 3--4(--5.5) cm, the outer greenish white to white or pink, the inner white; ovary with a few bracteoles (1--2 mm), hairs mostly lacking (except scant wool at base of fl.). Fr. narrowly ovoid, (2.5--)3.3--7.5 cm, red to lavender pink; seeds +- comma-shaped, (2--)3.5--4 mm, surface deeply pitted, dull, black.

Wet forest, 5--650 m; from both slopes, more or less throughout. The species, Mex.--Brazil; the variety, Nic.--Pan.

The thick, brittle cladodes, often with their lobe margins concave, and especially the very conspicuously pitted seeds, mark this species. I consider all CR material to be var. pittieri (F. A. C. Weber) Kimnach. One specimen (Bello 2363) has unusually small seeds, and another (Gomez et. al. 23806) is said to have diurnal fls. visited by Trigona bees.

Epiphyllum thomasianum (K. Schum.) Britton & Rose, Contr. U.S. Nat. Herb. 16: 259. 1913; Phyllocactus thomasianus K. Schum., Monatsschr. Kakteenk. 5: 6. 1895.; E. costaricense Britton & Rose, E. macrocarpum Back. E. macropterum sensu Britton & Rose.

Cladodes 23--95+ x 6--10 cm, the apex rounded to acute, the lamina drying dark gray to nearly black, sometimes brown along the edge; margins boldly crenate, the lobes mostly 35--55 x 5--10 mm, broadest at their middle, convex or rarely truncate. Bracts of areoles ca. 0.2--0.3 mm, the areoles with scant wool, sometimes with a few reddish hair-spines. Fls. trumpet- or funnel-shaped, 26--38 cm; tepals 9--14 cm, the outer rose colored, the inner white; ovary with a few bracteoles (ca. 0.5 mm), with wool and often a few hair-spines 6--10 mm. Fr. ovoid, ca. 10 cm, magenta; seeds pear-shaped, 3.2--3.5 mm, surface smooth, shiny, black.

Wet forest, 50--1300 m; Cordillera Tilaran and Llanura de San Carlos to Cordillera Talamanca (Atlantic slope), Osa Peninsula. Mex., Guat, Nic.--Pan., Ecu.

Perhaps most easily recognized in dried material by the nearly black lamina of the cladodes. In life, the lamina are unusually rubbery. Various authors have recognized two varieties, both presumably occuring in Costa Rica: var. thomasianum, lacking hair-spines, lamina margin green, tepals 12--13.5 cm, frs. smooth; var. costaricense, spine hairs present, lamina margin tan, tepals 9--10 cm, frs. ribbed. This species is often cultivated around San Jose.

Hylocereus

Climbing, viny epiphytes; stems 3-angled, usually with short, stiff spines on the aureoles, but none on the fls. or frs. Large fls.

Hylocereus calcaratus (F. A. C. Weber) Britton & Rose, Contr. U.S. Natl. Herb 12: 428. 1909; Cereus calcaratus F.A.C. Weber, Bull. Mus. Hist. Nat. 8: 458. 1902.

Stem wings 1.5--3.5 cm wide, thin, dark green in fresh material; margins scallop-lobed with bold protuberances near the areoles, the prouberances 1--2.5 cm from base to apex; areoles 3--5.5 cm apart, borne just above the protuberances, bearing dense wool and usually a few hair-spines 2--3 mm. Fls. funnel-shaped, 35--37 cm; tepals 10--15 cm, the outer greenish yellow, the inner creamy white; stigma lobes 18--21, not forked; ovary covered with large, broadly ovate, overlapping bracteoles, 0.5--2 cm, the lower ones covering a row of thin, flat hairs. Fruit and seeds unknown.

Wet forest, ?? m; Limon. Endemic.

I have a collection of a dead ringer for it from Cerro de La Muerte, 1750 m, live but never fertile.

Hylocereus costaricensis (F. A. C. Weber) Britton & Rose, Contr. U.S. Natl. Herb. 12: 428. 1909; Cereus trigonus var. costaricensis F.A.C. Weber, Bull. Mus. Hist. Nat. (Paris) 8: 457. 1902.

Stem wings 1--3 cm wide, usually very thick, grayish green, +- glaucous in fresh material; margins shallowly scallop-lobed, the lobes (areole to areole) 2--3.5 x 0.1--0.2 cm, the areoles bearing dense, short wool and 3--6 (9) short, dark spines 2--4 mm. Fls. funnel-shaped, 22--30 cm; tepals 11--15 cm, the outer greenish yellow, the inner white; stigma lobes ca. 12, not forked; ovary covered with large, broadly to narrowly triangular, overlapping bracteoles, 0.5--3 cm. Fruit broadly ovate to globose, bright magenta; seeds pear-shaped, black.

Dry forest and coastal areas, 0--1400 m; Pac. slope from Guanacaste to Rio Parrita valley. Nic., CR

Hylocereus monacanthus (Lem.) Britton & Rose, Publ. Carnegie Inst. Wash. 248(2): 190. 1920; Cereus monacanthus Lem., Hort. Univ. 6: 60. 1845.

Stem wings 0.5--2 cm wide, thickish, pale green to somewhat glaucous in fresh material; margins shallowly scallop-lobed, the lobes 3.5--6 x 0.3--0.4 cm (concave between the areoles in dry material), the areoles with scant, short wool (young stems) and 1 or 2(3), short, ambar spines 1.5--2 mm. Fls. funnel-shaped, 20--28 cm; tepals 9--13 cm, the outer yellow with red margins, the inner white but shaded pink towareds base; stigma lobes ca. 15, forked; ovary with several large, broadly triangular, but distant bracteoles, 0.5--1 cm. Fruit and seeds unknown?

Dry forest, riverside, 50--700 m; Pacific slope, Valle de El General--Rio Terraba drainage. CR and Pan. Aguilar & Quesada 2085, Hammel 18213, Solomon 19314.

In CR known only from the southern part of the country, Pacific slope. Not only is the name tentative, the separation of this from what we call H. costaricensis in CR is also hypothetical, supported primarily by distribution, number and size of spines, and size of the ovary bracteoles, branching of the stigma lobes.

Hylocereus stenopterus (F.A.C. Weber) Britton & Rose, Contr. U.S. Natl. Herb. 12: 429. 1909; Cereus stenopterus F.A.C. Weber, Bull. Mus. Hist. Nat. (Paris) 8: 458. 1902.

Stem wings 1.5--2 cm wide, thin, pale green in fresh material; margins shallowly scallop-lobed, the lobes 3.5--4.5 x 0.3--0.4 cm (concave in dry material), the areoles bearing dense, short wool and (0)1--3, short, tan spines ca. 2 mm. Fls. funnel-shaped, ca. 13 cm; tepals 7--9 cm, the outer reddish purple, the inner pink to purplish red, nearly white along the margin; stigma lobes 14--18, forked; ovary covered with large, broadly ovate, overlapping bracteoles. Fruit globose, 7 cm, yellowish green (immature); seeds pear-shaped, 2.5--3 mm, the surface smooth to shallowly pitted, black.

Wet forest, 15--1000 m; Atl. slope, Llanuras de San Carlos and Tortuguero to Cord. Talamanca. Endemic.

This species is distinctive for its relatively small and red fls. The stems are 3 angled, the wings thin, and sometimes without spines. What's the diff. between this and H. monacanthus in sterile cond.? (Alfredo Brade 110 & 123, CR; M.M Chavarria 576, CR, INB)

Melocactus

Melocactus curvispinus Pfeiff., Enum. Diagn. Cact. 46. 1837. M. maxonii (Rose) Gurke, M. maxonii (Rose) Gurke var. sanctae-rosae L.D. Gómez, M. ruestii K. Schum., M. ruestii K. Schum. subsp. sanctae-rosae (L.D. Gómez) Elizondo.

Terrestrial, short-stemmed, globose to cylindric to ca. 30 cm tall, the stems with 10--16 ribs, the ribs ca. 10--27 mm wide.

Rare, Peninsula de Santa Elena and nearby. A small "barrel" cactus with reddish fls.

Nopalxochia

Nopalxochia horichii Kimnach, Cact. Succ. J. (Los Angeles) 56: 6. 1984. Disocactus kimnachii G.D. Rowley (based on same type as above)

Laminate portion of stems (cladodes) rather thick (6--10 mm), flattened or 3-angled, 2--4 cm wide, the apex acute or rounded, the lamina drying dull, light grayish green; margins serrate lobed, the lobes (areole to areole x base to apex) 15--30 x 1--3 mm, broadest near the top, the areoles with abundant and conspicuous wool and several (1--5+) slender but stiff spines, ca. 3--5 mm. Fls. +- funnel-shaped, said to be 16.5--17 cm long, but with a rather long tube, and in supposed clonotype material only just over half that size (10 cm); tepals 5--6 cm pink; ovary bracteoles ca. 1 mm with conspicuous, tan wool and stiff spines 4--6(--9) mm. Frs. +- globose, 2--3 cm, red with pink pulp, the wool and spines persistent; seeds pear-shaped, 1.5--1.7 mm, conspicuously rugose-pitted, shiny black.

Wet forest, 1400 m. Cordillera Volcanica Central.

A nightmare! I may treat this as just a cultivated, pot ornamental, only mention it under the family.

Opuntia

Terrestrial with obovate usually spiny "pads;" fls. with no tube, yellow or red.

Opuntia brasiliensis (Wild.) Haw., Suppl. Pl. Succ. 79. 1819; Cactus brasiliensis Willd., Enum. Pl. 33. 1813.

Stems erect to

This is the tall-trunked, cultivated one with small, thin pads, cylindrical trunk and side branches; yellow fls. Perhaps should be treated. It persists in fence rows around the Valle Central.

Opuntia cochenillifera (L.) Mill., Gard. Dict. (ed. 8) No. 6. 1768. Cactus cochenillifer L., Sp. Pl. 468. 1753. Nopalea cochenillifera (L.) Salm-Dyck.

We shall use Opuntia as for Flora Nicaraguensis. Cultivated but should be treated.

Opuntia ficus-indica (L.) Mill., Gard. Dict. (ed. 8) Opuntia n. 2. 1768; Cactus ficus-indica Sp. Pl., 1: 468. 1753.

This is commonly cultivated, probably should be treated.

Opuntia guatemalensis Britton & Rose, Cact. 1: 218, f. 285. 1919.

Dry forest, coastal thickets, 0--500 m; Pac. slope, Guanacaste and Valle Central.

Lots of collections of this one. It is a small, usually sprawling plant with variable length spines, usually white with dark (or green in life) tips.

Opuntia lutea Opuntia lutea (Rose) D. R. Hunt, Cact. Consensus Init. 4: 6. 1997; Nopalea lutea Rose, Contr. U.S. Natl. Herb. 12:405. 1909.

Dry forest, 100 m; known only from Palo Verde (Tempisque valley). Guat. to CR. U. Chavarria 827, CR, INB

Peniocereus

Peniocereus hirschtianus (K. Schum.) Hunt, Bradleya 9: 90. 1991; Cereus hirschtianus K. Schum., Gesamtbeschr. Kakt. 130. 1897. Nyctocereus hirschtianus (K. Schum.) Britt. & Rose.

Dry forest, coastal thickets, near sea level; Guanacaste, especially Santa Elena Peninsula.

I've seen it in dune vegetation along the beach, trailing on the ground, cylindrical and ribbed densely covered with short spines. Rarely cultivated.

Pereskia

Terrestrial with spines and obvious, real leaves.

Pereskia grandifolia Haw., Suppl. Pl. Succ. 85. 1819.

Cultivated but should be treated in the Manual. Pink flowers.

Pereskia lychnidiflora DC., Prodr. 3: 475. mar. 1828. P. nicoyana F.A.C. Weber

Orange flowers.

Dry forest thickets, 8--75 m; Guanacaste, Valle del Tempisque. Mex. to CR. Ulises Chavarria 788

Rhipsalis

Pendulous, spineless epiphytes, the stems terete, angled or flat (elsewhere), polychotomous branching unique?; fls. very small (<1 cm?), the ovary lacking bracteoles; frs. globose; seeds elongate ovoid.

Rhipsalis simmleri P. Beauv. (Bull. Herb. Boissier, ser. 2, 7: 136. 1907) was reported by Barthlott & Taylor (1995) as a synonym of R. cereuscula Haworth. It was based on material said to have arrived (Paris) among a shipment of bromeliads from Costa Rican, but undoubtedly a horticultural mix-up. The species is South American, mostly from dry areas of SE Brazil, Bolivia, Paraguay, Uruguay, has the ultimate stem segments angled, often with hair-spines, the flowers pendulous. The plants that Horich claimed to have collected near Caño Negro, and which he said were possibly this species, are R. teres (see Barthlott & Taylor, Bradleya 13: 65.1995), and certainly neither native nor introduced (except by Horich himself!) to Costa Rica.

Rhipsalis baccifera (J.S. Muell.) Stearn, Cact. J. (Croydon) 7(4): 107. 1939; Cassytha baccifera J.S. Muell. Ill. Syst. Sex. Linnaei [t. 2a]. 1771. Rhipsalis cassutha Gaertn.

Wet forest, 10--850 m; Atl. slope more or less throughout, Pac. slope from Cord. Guanacaste, Carara Reserve, Valle de El General and Osa Peninsula. Common, especially on the Atl.

Long, cylindrical drooping stems, small white fls. & frs.

Rhipsalis micrantha (Kunth) DC., Prodr. 3: 476. 1828; Cactus micrantha Kunth, Nov. Gen. Sp. 6: 65. 1823. R. tonduzii F. A. C. Weber; R. wercklei Berg.

Seasonal wet or moist forest, 800--1400 m; Pacifico Central, Fila Diamante.

Angled stems, small, white, round frs.; most easily found in moist forest, e.g., near El Rodeo.

Selenicereus

Climbing, viny, sometimes closely clinging or loosely hanging epiphytes; stems spiny or not, cylindrical and many ribbed, or broadly several-winged; fls. large with long tube, at least the fr. spiny.

Selenicereus testudo (Karw.) Buxb., Kakteen 1(VI): CIIa. 1965; Cereus testudo Karw., Akademie der Wissenschaften 2: 682. 1837; Deamia testudo (Karw.) Britton & Rose. Selenicereus miravallenis (F. A. C. Weber) Britton & Rose.

Plants climbing and tightly clinging on rocks and trees to several meters, wrapped around branhes like a snake; stems strongly 3--7 angled, the wings 1--7 cm wide, rather thin, all of them often drawn down towards the substrate and forming chambers inhabited by roots and ants, green, not glaucous in fresh material, in dry material often with shiny and exfoliating cuticle; margins shallowly crenate, the lobes 1--3 x (0--)0.1--0.3 cm, the areoles with dense wool and numerous tan to reddish brown spines 5--15+ mm, often also with hair-spines, the spines and spine-hairs often bearing abundant, minute (<0.1 mm) hairs or pustules. Fls. 20--27 cm; tepals 8--10 cm, white to cream-colored; ovary bracteoles minute, subtending a dense ball of tan wool and numerous orange to reddish brown spine-hairs to 20+ mm, often also with short spines. Frs. globose, ca. 5? cm, bright red, the pulp white, the wool and stiff spines persistent; seeds somewhat elongate, pear-shaped, 1.8--3 mm, smooth, shiny black.

Hugs tree branches, usually 5--7 angled, broad, thin wings, longish spines, large globose, spiny, red fruits with white pulp.

Dry forest, 0--250 m; Guanacaste. Solomon 603 (CR).

Selenicereus wercklei (F.A.C. Weber) Britton & Rose, Cact. 2: 208, f. 288, 289. 1920; Bull. Mus. Hist. Nat. (Paris) 8: 460. 1902. No type designated.

Stems cylindrical, 5--15 mm diam., 6--12 ribbed; areoles 4--7 cm apart, with a small tuft of wool, but no spines, except rarely on young stems. Fls. 16--23 cm; tepals 5--7 cm, the outer yellowish green, the inner white with bright pink at base; ovary with numerous, small, bracteoles but these mostly obsured by the dense, tan wool and redish brown spines, 1--8 mm. Frs. ovoid, 5--6 cm (immature), yellowish green, the wool and stiff spines persistent; seeds pear-shaped, 1.5--? mm, ?, ? ?.

Moist to wet forest, 600--1100 m; Cord. Guanacaste--Cord. Tilaran, then disjunct to Rio Terraba canyon. Endemic.

This species is well known, with numerous specimens at INB and CR. Cylindrical, ribbed stems, no spines except on flowers. (Hammel 17810). Dried sterile material may be difficult to distinguish from Weberocereus biolleyi, but which tends to have lighter gray drying stems and no more than 3- or 4-ribbed, except falsely on drying.

Stenocereus

Stenocereus aragonii (F.A.C. Weber) Buxb., Bot. Stud. 12: 99. 1961; Cereus aragonii F.A.C. Weber, Bull. Mus. Hist. Nat. (Paris) 8: 456. 1902.

The columnar one with the glaucous herringbone pattern at the tip.

Dry forest, coastal hills and river canyons, 10--50 (800) m; Guanacaste, Valley Central (Rio Grande y Virilla canyons), then disjunct S to Rio Terraba canyon. Endemic? Chavarria 789.

Weberocereus

Pendulous or viny and climbing epiphytes with all manner of stems, with or without spines; fls. medium sized, short, usually tuberculate (ovary) tube, sometimes with spines on the fr. The presence or absence of spines on the stems is variable in nearly all species, some usually having spines, others usually not. No CR species is totally without stem spines. Seeds markedly pitted in all CR species except W. tonduzii.

Weberocereus biolleyi (F.A.C. Weber) Britton & Rose, Contr. U.S. Natl. Herb. 12: 431. 1909; Rhipsalis biolleyi F.A.C. Weber, Bull. Mus. Hist. Nat. (Paris) 8: 467. 1902.

Stems cylindrical, 2--7 mm diam., smooth to obscurely 3 or 4 angled; areoles 2--4.5(--7) cm apart, with mostly obscure wool, occasionally a few, flimsy hair-spines, but seldom with stiff spines. Fls. ca. 4--5(--6) cm; tepals 1.5--2.5 cm, the outer green to pink, the inner white to pink; ovary with numerous, small, broadly triangular bracteoles (ca. 0.5 mm) raised on conspicuous bumps and bearing dense wool and orangish hair-spines 5--10+ mm. Frs. ovoid, 1.5--2.5 cm, reddish purple with purple pulp, wool and hair-spines persistent; seeds pear-shaped, 1.2--1.5 mm, surface conspicuously pitted, dull black.

Wet forest, 2--200 (--1100) m; Atl. slope, from Llanuras de Tortuguero at least to Playa Bananito (S of Limon). CR, Pan.

Slender, smoothly cylindrical, glabrous stems, small pink fls., common on the Atl. lowlands. N.B. are plants with triangular stems just a form of this, or a distinct species--see W. panamensis. Also, occasional specimens have spines and may therefore be confused with W. trichophorus.

Weberocereus bradei (Britton & Rose) G.D. Rowley, Rep. Pl. Succ. 23(1972): 10. 1972 [1974]; Eccremocactus bradei Britton & Rose, Contr. U.S. Natl. Herb. 16: 262, t. 83. 1913; Epiphyllum bradei (Britton & Rose) Standl.

Stems flat and thick (ca. 5 mm), the cladodes 13--60+ x 5--17 cm, the apex rounded, the lamina drying dull, yellowish tan; margins shallowly crenate, the lobes 2.5--4.5 x ca. 3 cm, broadest towards their middle. Areoles with dense wool and often with 4--7 stiff orangish spines up to 1 cm. Fls. 5--8 cm; tepals 2.5--3.5 cm, the outer pink, the inner white to; ovary with few, small, narrowly triangular bracteoles (0.5--3 mm) raised on ridges and subtending dense wool, sometimes with a few hair-spines. Frs. ovoid, 3.5 cm, browinsh red with reddish pulp; seeds pear-shaped, ca. 1.3 mm, surface conspicuously pitted, dull black.

Seasonal wet forest, 20--850 m; Pac. slope only from Pacifico Central (Carara--Fila Bustamante region) and the Osa Peninsua. Endemic.

By virtue of its large, thick, obovoid "pads" (even sometimes with spines on the margin) it looks like an epiphytic Opuntia.

Weberocereus imitans (Kimnach & Hutchison) Buxb., Succulenta (Amsterdam) 57(6): 125. 1978; Werckleocereus imitans Kimnach & Hutchison, Cact. Succ. J. (Los Angeles) 28(5): 154. 1956.

Stems flat, relatively thin (< 2mm), the cladodes 35--60 x 6.5--15 cm, the apex +- acute, the lamina drying dull, yellowish tan; margins very deeply lobed nearly to the midrib, the lobes 1.5--2.5 cm wide (top to bottom), aimed upwards. Areoles 2.5--5 cm apart, with mostly obscure wool, occasionally a few, flimsy hair-spines, but seldom with stiff spines. Fls. 5.5--7 cm; tepals ca. 2.5 cm, the outer yellow with a reddish center line, the inner cream; ovary with few, small, broadly triangular bracteoles (< 0.5 mm) raised on obscure bumps and bearing dense wool and slender, orange spines 3--5 mm. Frs. ovoid, ca. 3.5 cm, green to red with white pulp, the spines and wool of the ovary persistent, spines to 10+ mm; seeds pear-shaped, 1.3--1.4 mm, surface conspicuously pitted, dull black.

Seasonal wet forest, 20--900 m; Pac. slope, only from S part of the country, mostly from the Osa Peninsula and nearby, with a few collections from Valle de El General. Endemic.

Easy to distinguish among all native CR, flat-stemmed species for its very deeply lobed stems, short, white fls., no spines (except on frs.). Known only from southern Pac. slope.

?Weberocereus panamensis Britton & Rose.

Stems sharply 2--4 winged, 7--17 mm wide; areoles 3--4 cm apart, with mostly obscure wool, lacking both hair-spines and spines. Fls. ca. 4--5(--6)? cm; tepals 1.5--2.5? cm, white?; ovary with numerous, small, broadly triangular bracteoles (ca. 0.5 mm) raised on conspicuous bumps and bearing dense wool and orangish hair-spines 5--10+ mm. Frs. ovoid, ca. 3 cm, reddish purple with purple flesh, wool and hair-spines persistent; seeds pear-shaped, 1.2--1.4 mm, surface conspicuously pitted, dull black.

Wet forest, 1000? m; Cord. Guanacaste, road to Upala.

This is most likely just a form of W. biolleyi. I've seen no fls., fl. measurements here are taken from those of W. biolleyi.

Weberocereus tonduzii (F.A.C. Weber) G.D. Rowley, Natl. Cact. Succ. J. 37(2): 46. 1982; Cereus tonduzii F.A.C. Weber, Bull. Mus. Hist. Nat. (Paris) 8: 459. 1902.

Plants climbing on rocks and trees to several meters; stems strongly 3 angled, the wings 1--2.5 cm wide, rather thick, green, not glaucous in fresh material; margins shallowly crenate, the lobes 2--2.5 x 0.2--0.3 cm, the areoles with scant wool and usually with 1--5 short, dark spines ca. 1 mm. Fls. ca. 6--7(--8) cm; tepals 1.5--2.5 cm, the outer greenish yellow to pink, the inner cream; ovary bracteoles minute or lacking, slightly raised on low bumps and bearing a dense ball of nearly black wool and reddish brown spines to 5 mm. Frs. globose, ca. 4 cm, yellow with cream-colored pulp, the wool and spines persistent; seeds pear-shaped, 1.5--2.2 mm, smooth, shiny black.

Wet forest, 100--2500 m; Pac. slope, Cord. Volcanica Central (Grecia) and Cord. Talamanca. Endemic. Gina Umana 253,

Rare species from rather high elevation. The unpitted seeds are unusual for the genus, perhaps belongs with Selenicereus?

Weberocereus trichophorus H. Johnson & Kimnach, Cact. Succ. J. (Los Angeles) 35: 205. 1963.

Plants with abundant adventitious roots, climbing high in trees; stems cylindrical, 8--12 mm diam., smooth to obscurely 5--7 angled; areoles (spine clusters) ca. 1.5 cm apart, with copious tan wool, and many hair-spines (5--20 mm) and stiff spines (3--12 mm). Fls. ca. 4--6 cm; tepals 1.5--2.5 cm, the outer pink ("rose red"), the inner white; ovary with numerous, +- ligulate bracteoles (0.5--1 mm) raised on low bumps and bearing dense wool, but all obscured by the abundant, orangish hair-spines ca. 10 mm. Frs. ovoid, 2--3 cm, reddish, the pulp reddish purple, the hairs persistent, some of them as stiff spines; seeds pear-shaped, 1.1--1.3(--1.7) mm, surface conspicuously pitted, dull black.

Wet forest, 1--100 m; Atl. slope, Puerto Viejo de Limon to Sixaola. Endemic.

Easy to distinguish for its slender, cylindrical stems with spines and rusty, woolly hairs. Known only from the Atl. coast S of Cahuita. It is surely in Pan., and I've seen sterile material from Nic. The parenthetical and large seed measurement is from Johnson and Kimnach's original description of a greenhouse plant; the smaller size range being that found in fruits of field-collected, herbarium material, Hammel 18140. As mentioned by Johnson and Kimnach, the earlier name Cereus estrellensis F.A.C. Weber ex Werckle, no type specimen known, may refer to this species. It is described as a climbing, cylindrical and spiny stemmed cactus from the same region where W. trichophorus is the only cactus known that could fit that description. Good enough reason to bring that name out of limbo? Need to get Werkle's mention of this translated.

Weberocereus tunilla (F.A.C. Weber) Britton & Rose, Contr. U.S. Natl. Herb. 12: 432. 1909; Cereus tunilla F.A.C. Weber, Bull. Mus. Hist. Nat. (Paris) 8: 460. 1902. Cereus gonzalezii F. A. C. Weber. No type designated, original description probably from live material from greenhouse in Paris, sent to Weber by way of Werckle. Type (Werckle s.n., Tablon near Cartago, 1400 m; ca. 1901) would be at P if such exists.

Stems (2--)4(--6) angled, 6--23 mm diam., the areoles 2--4(--6) cm apart, with dense wool and numerous stiff spines (3--5 mm). Fls. 6--7 cm; tepals 1.3--2.7 cm, the outer pinkish ambar, the inner pink; ovary with numerous, triangular bracteoles (ca. 1--2 mm) raised on conspicuous bumps and bearing dense wool and white hair-spines 2--8 mm. Frs. ovoid (oblong), 4.5 cm, light pink, the pulp dark red, wool, hair-spines and stiff spines persistent; seeds pear-shaped, 1.5--2 mm, surface conspicuously pitted, dull black.

Wet forest, 1400 m; near Cartago. Endemic.

I've seen no material of this: the description here is modified directly from Kimnach (1968. C & S J, 40(3)113--115). Since his measurements were certainly taken from live material and those here of other species from dry material, size differences especially of stem width and fl. length may be exagertated.

Schombergara truncatus or whatever the common cult. Species (Christmas Cactus) is, should be mentioned only under the family description.

Unknown species published from CR

Cereus estrellensis F. A. C. Weber Monatsschr. Kakteenk. 15: 167. 1905.

According to Britton & Rose "the stems are 6-angled; the flowers are small, rosy to salmon-colored, and nocturnal....may belong to our genus Weberocereus" They say that Wercklé reports it as similar to Cereus nycticaullus (=Selenicereus pteranthus) but weaker and more spiny. By this and the location (Valle de Estrella, I presume) it might very well be an older name for W. trichophorus. Or a synonym of W. biolleyi. Johnson and Kimnach also mentioned this problem when they described W. trichophorus.

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