Garrett E. Crow
Lloyd, F. E. 1942. The Carnivorous Plants. Chronica Botanica. Waltham, Massachusetts, USA. Taylor, P. 1975. Lentibulariaceae. In G. Harling and N. B. Sparre (eds.) Flora of Equador. Opera Bot. ser. B, 4: 9-21.
Taylor, P. 1976. Flora of Panama. Family 176. Lentibulariaceae. Ann. Missouri Bot. Gard. 63: 565-580.
4 gen., ca. 232 spp.; nearly cosmopolitan; 1 gen., 10 spp. in CR.
Perennial or annual; aquatic, amphibious, terrestrial or epiphytic herbs; carnivorous. Leaves entire to much divided, bearing sticky glands (Pinguicula) or bladder-like traps (Genlisea, Polypompholyx, Utricularia). Stems short, with leaves in basal rosette, or elongate, bearing dissected leaves. Inflorescences racemose, bracteate, or flowers solitary, scapose. Flowers bisexual, zygomorphic, subtended by a pair of bracteoles. Sepals 2-, 4- or 5-lobed. Petals 5, connate at base, bilabiate, lower lip saccate or spurred. Stamens 2, epipetalous at base of corolla tube, anthers 1-celled. Pistil with 2 carpels, ovary superior, placentation free-central (or basal). Fruit a capsule, circumscissile, or dehiscing irregularly by 2 or 4 valves. Seeds usually numerous, small.
Bladderwort
Crow, G. E. 1992[1995]. The genus Utricularia (Lentibulariaceae) in Costa Rica. Brenesia 38: 1-18. Taylor, P. 1989. The genus Utricularia--a taxonomic monograph. Kew Bull. Add. Ser. XIV: 1-724.
Ca. 214 spp., nearly cosmopolitan, tropical to cold temperate; 10 spp. reported for CR.
Epiphytic, terrestrial, and amphibious or submersed aquatic herbs, carnivorous. Roots absent, usually anchored by rhizoids, stolons usually present. Aquatic species perennating by turions. Leaves of epiphytic species petiolate, usually arising from tubers, with bladders (carnivorous traps) borne on separate non-photosynthetic branch systems. Aquatic species with lateral leaf-like photosynthetic branch systems, bearing bladders. Flowers zygomorphic, borne on scapes, solitary or in racemes. Calyx 2-lobed. Corolla 2-lipped, lower lip usually 3-lobed, mostly with a distinct palate, spurred. Stamens 2, epipetalous. Fruit a 2-valved capsule.
Although the photosynthetic organs are actually modified stems (McIntyre and Chrysler, 1943), they are described as leaves for this treatment.
Utricularia subulata, a nearly pantropical plant of wet sandy/peaty savannas should be expected in CR; it has been collected in northeastern Nicaragua and in Panamá. Several other taxa might be expected to occur in Costa Rica. The following taxa have been reported for Panamá, but not reported for Costa Rica: U. asplundii, U. lloydii, U. panamensis. Taxa which have been collected in Nicaragua (savannas), but not reported for Costa Rica include: U. olivacea, U. tricophylla, U. benjaminiana, U. erectifolia, U. guyanensis, U. hispida, U. juncea.
2. Calyx lobes not alike, lower lobe shorter than upper; spur of corolla 2-3 times as long as the lower lip, diverging laterally, nearly straight; tubers absent (Sect. Foliosa)...U. amethystina
2. Calyx lobes alike, equal; spur of corolla shorter than or equalling lower lip, or when greatly exceeding, then strongly curved; tubers present, ellipsoid to nearly globose (Sect. Orchidoides).
3. Corolla 1-1.5 cm long; lower lip of corolla more or less deeply 3-lobed, spur much longer than lower lip; calyx 0.3-1 cm long; leaves 0.7-2(4.5) cm long...U. jamesoniana
3. Corolla 3-5 cm long; lower lip of corolla entire or shallowly lobed, spur as long as or shorter than lower lip (seldom slightly exceeding); calyx (1)1.5-3.5 cm long; leaves 3-15 cm long.
4. Corolla (both surfaces and all margins) densely covered with slender stipitate glands...U. endresii
4. Corolla essentially glabrous, or with a few marginal stipitate glands.
5. Scales and bracts (1)1.5-2(3) cm long, curved; corolla spur apex not deflexed; upper lip of corolla shorter than and narrower to equalling calyx; leaves narrowly elliptic...U. unifolia
5. Scales and bracts 0.5-0.7(1) cm long, straight; corolla spur double curved, apex deflexed; upper lip of corolla as long as and slightly wider than calyx; leaves broadly elliptic to broadly oblanceolate...U. praetermissa
1. Plants aquatic, submersed, or in wetlands, appearing terrestrial, but anchored in wet substrate and apparent only when flowering; leaves divided into filiform, branch-like segments (slender oblanceolate vegetative lamina sometimes present in U. pusilla); bracts present, bracteoles absent.
6. Plants with leaf branches whorled; bladders borne at tips of leaf segments; flowers light purple, lobes of lower lip of corolla 3, lateral lobes saccate (Sect. Vesiculina)...U. purpurea
6. Plants with leaf branches chiefly alternate; bladders borne scattered on branch segments; flowers yellow, or if pink to light purple, then lobes of lower lip of corolla 2, not saccate (U. hydrocarpa).
7. Plants subaquatic, appearing terrestrial, with rhizoids anchored in wet substrate; bracts peltate; spur of corolla twice as long as the lower lip; lamina frequently present, narrowly oblanceolate, mostly 1 mm wide, 1-nerved; ultimate branch segments never laterally setulose (Sect. Setiscapella)...U. pusilla
7. Plants submersed aquatics (sometimes stranded on mud as water table drops); bracts basifixed; spur of corolla shorter than, equalling or slightly longer than the lower lip; lamina absent; ultimate branch segments laterally setulose (Sect. Utricularia).
8. Corolla pink to light purple; bladders dimorphic, smaller traps about half the size of the larger traps...U. hydrocarpa
8. Corolla yellow; bladders alike.
9. Flowers usually few, (1)2-3(6, very rarely to 12); upper corolla lip larger than the lower; stolons terete; vegetative branches essentially alike, dichotomously branched (mostly up to 4 times, occassional to 8 times), ultimate segments short, not filamentous; capsule dehiscent into 2 valves...U. gibba
9. Flowers usually several-many (3)8-20); upper corolla lip smaller than the lower; stolons flattened; vegetative branches usually dimorphic (branches essentially bearing bladders/branches essentially vegetative, bearing few bladders, alternately branched (with multiple pinnae), ultimate segments of vegetative branches long, exceedingly fine, filamentous; capsule indehiscent...U. foliosa
Utricularia amethystina Salzmann ex A. St. Hil. & Girard, Compte Rend. Hebd. Séances Acad. Sci., D 7:870. 1838.
Annual or perennial. Leaves very small, 2-6 in a rosette, lamina (narrowly to) broadly obovate to nearly orbicular, up to 6 mm wide. Tubers absent, Flowers (1)2-6(10). Calyx lobes unequal, + densely glandular pubescent. Corolla white to lavender to light purple, with a yellow spot at base of lower lip, lower lip shallowly 3-lobed; spur strongly laterally projecting, 2-3 times the length of the lower lip. Capsule 1.5-2.5 mm diameter.
Terrestrial; wet sandy savannas, peaty swamps, damp soil among rocks and along streams. San Miguel Oeste, Naranjo (Herrera 24, MO); La Palma de San Ramón (Brenes 642, F). Fl. Oct, Nov. Florida, Mex.—Peru, Bra., and Bol.
Utricularia endresii H.G. Reichb., Gard. Chron. 1874(2): 582. 1874.
Perennial. Leaves 1-3, 6-20 cm long, lamina membranous to thinly coriaceous (often quite membranous when dry), narrowly oblanceolate to narrowly oblong-elliptic to linear-lanceolate, 5-12 cm long, 0.5-1.5 cm wide; all vegetative parts covered with globose or cylindrical glands. Bracts mostly 5-6 mm long, narrowly linear. Flowers 1-4(6), large. Corolla 3-4(5) cm long, lavender to violet, with yellow blotch at base of lower lip, both surfaces and margins densely covered with stipitate glands, lower lip nearly entire, wider than long; spur subulate, apex acute, curved upwards, about as long as lower lip. Capsule ovoid, ca. 8 mm long.
Epiphytic or terrestrial; on mossy tree trunks and wet banks in montane rain forest. Mainly 350-850 m elevation. San Ramón (Brenes 33583, F), Térraba (León 1100, CR), Faldas del Cerro Nara, ca. Esquipulas y Tarrazú (Gómez-Laurito, López & Gómez 11606, CR, F. Fl. Jul.—Jan. CR—Pan., Col. and Ecu. Grows in drier habitats than the epiphytic species; looses its leaves in dry season.
Utricularia foliosa L., Sp. Pl. 1: 18. 1753. U. mixta Barnhart, Mem. Torrey Bot. Club 23: 105. 1920.
Perennial, submersed aquatic. Stolons long, robust, distinctly flattened, 1-4 mm wide, very mucilaginous, often branched. Leaves (modified branches) numerous, alternate, with a single long primary rachis (often 20-30(45) cm long) bearing numerous pinnate, secondary leaf segments (pinnae) with highly dissected, extremely fine capillary segments. Leaves somewhat dimorphic with some bearing numerous traps, others bearing few or none. Traps 1-2 mm long. Inflorescence erect, emergent. Bracts basifixed. Flowers (3)7-15(20). Corolla 0.8-1.5 cm long, yellow, palate of lower lip conspicuous, sometimes with reddish streaks, lower lip shallowly 2-lobed; spur narrowly conical, straight, somewhat shorter than or equalling lower lip. Capsule globose, up to 8 mm diameter.
Submersed; quiet or slowly moving water, lakes, shallow pools, rivers, marshes, and ditches. Low elevations. Llanura Los Guatusos (Gómez-Laurito 10853, CR), Tilaran (Ocampo 905, CR), Valle de Paraíso de Cartago (Soto s.n., CR), Guanacaste lowlands, Valle del Tempisque (Crow & Rivera 6281, CR, F, MO, NHA). Fl. throughout year. Widespread, trop. Amer., Afr. and Madagascar; se. USA, Mex.—Pan., S. to Bol. and Argentina, Antilles. This species is particularly distinctive with its flattened stolons and extremely fine ultimate branches, which readily collapse when removed from water, and by its robust, many-flowered inflorescence. Taylor (1989) notes that the fruits are berry-like capsules which float, but that the seeds sink immediately.
Utricularia gibba L., Sp. Pl. 1: 18. 1753. U. obtusa Sw., Prodr. Veg. Ind. Occ. 14. 1788.
Perennial or annual, submersed aquatic. Stolons numerous, much branched, often forming tangled mats. Leaves (modified stems) numerous, 0.5-1.5 cm long, alternate, highly dissected, primary segments usually 2, dichotomously branched, segments capillary. Traps relatively few, small, 1-1.5 mm long. Inflorescence erect, emergent, or sometimes bearing submersed cleistogamous flowers. Bracts basifixed. Flowers mostly 1-2, small. Corolla (3)4-6 mm long (measured from base of corolla to tip of lower lip), yellow, often with reddish streak on the palate of lower lip, upper lip obscurely (sometimes distinctly) 3-lobed, lower lip slightly smaller, entire; spur more or less narrowly conical, with apex obtuse, slightly shorter than to exceeding lower lip. Capsule globose, 2-3 mm diameter.
Shallow still waters of ponds, swamps, marshes, and pools; submersed or stranded on muddy substrate after water level drops. Lowlands, sea level to ca. 1000 m elevation. Llanura Los Guatusos (Crow & Rivera 7602, CR, F, MO, NHA), Laguna Fraijanes, Cordillera Central (Bumby 338, CR, F) Valle de Turrialba (Bumby 327, CR), Paraíso (Bumby 313, CR, MO), Valle del Tempisque (Crow & Rivera 5970, CR, F, MO, NHA), Hatillo, Nicoya Pen. (Crow & Rivera 6880, CR, F, MO, NHA), Barranca (Crow & Rivera 6014, CR, MO, NHA), Buenos Aires, Valle de El General (Crow 8971, CR, INBIO, MO, NHA), San Vito (Crow & Rivera 7575a (NHA), near Cañas Gordas (Barringer & Gómez 16687 (F), Valle de Parrita (Crow 7585 (CR, F, MO, NHA, RSA). Fl. most of year. Widespread, e. Can. and e. U.S.A. S. to Florida and Tex., Mex.—Pan., S. to Chile and Argentina; pantropical. Taylor (1975) previously treated material from this region as U. obtusa, however there is a great deal of variability in this complex and he has since placed U. obtusa Sw. in synonomy under U. gibba L. Taylor (1989) also includes the larger-flowered U. biflora Lam. in synonomy under U. gibba, however, all the material examined from Costa Rica has small flowers, typically 4-6 mm long. Flowering typically occurs when water is very shallow; plants in deeper water are usually vegetative, unless supported on floating mats of living or dead vegetation. Plants stranded on mud, flowering profusely, are frequently collected without the vegetative parts, which remain in the mud. Hence, these specimens are sometimes misdetermined as U. subulata. The latter species, however, has peltate bracts.
Utricularia hydrocarpa Vahl, Enum. 1: 200. 1804.
Perennial or annual, submersed aquatic. Stolons sparsely branched. Leaves (modified stems) numerous, 1.5-2.5 cm long, alternate, highly dissected, primary segments 2, repeatedly dichotomously branched, segments capillary. Traps relatively numerous, conspicuous (often black), dimorphic, the smaller traps ca. 1 mm long, larger 2-3 mm long. Inflorescence erect, emergent, with lowermost flower very close to the base of peduncle, submersed, strongly deflexed, cleistogamous. Bracts long, basifixed, 2.5-3(3.5) long. Flowers 3-7 (often 1 open at a time). Corolla 5-7(10) mm long (measured from base of corolla to tip of lower lip), pink to light purple, with yellow blotch at base of lower lip, lower lip wider than long, 2-lobed; spur conical, apex obtuse to subacute, shorter than the lower lip. Capsule globose, 2-3 mm diameter.
Submersed; lakes, ponds and shallow pools; elevations up to 700 m. Llanura Los Guatusos (Crow & Rivera 7599, CR, F, MO, NHA, RSA), Guanacaste lowlands near P. N. Santa Rosa (Crow 6229, CR, F, MO, NHA), La Cruz (Gómez 18955 (M0), Valle de El General, Buenos Aires (Crow 6174, CR, F, MO, NHA). Fl. Aug.—Dec. Mex., C.Am. S. to Bol. and Par.
Utricularia jamesoniana Oliver, J. Linn. Soc. Bot. 4: 169. 1860. U. verapazensis Morong ex J.D. Smith, Bot. Gaz. 18: 230. 1893. U. schimperi Schenck, Jahrb. Wiss. Bot. 18: 230. 1887.
Perennial, tubers present. Plants small, delicate. Leaves 1-3, 0.6-2(4.5) cm long, coriaceous, lamina obovate to spatulate to sometimes narrowly elliptic, 6-10 mm (sometimes 20-30 mm) long, 2-5(10) mm wide, tapering to a petiole. Flowers 1-3(-5), small. Calyx broadly ovate. Corolla 1-1.5 cm long, white or tinged with purple, yellow or yellow-purple streaked blotch at base of lower lip, lower lip more or less deeply 3-lobed, the palate not prominent; spur long, greatly exceeding the lower lip (3-5 times as long), strongly curved or bent upwards toward apex; entire corolla and especially spur densely covered with stipitate glands or simple hairs. Capsule ca 0.3 mm long.
Epiphyte; on mossy tree trunks in open pastures and secondary growth cloud forest habitats. 800-2100 m elevation. La Palma de San Ramón (Brenes 4464, CR, F), Monteverde, Cordillera de Tilarán (Dryer 24B, CR), Valle de Río Peñas Blancas (Haber ex Bello & Cruz 6100 (MO), Tapantí (Grayum 3966 CR, MO), Casa de Tejas, above Río Gato (Lent 3708, CR), Vara Blanca de Sarapiquí, Central Cordillera (Skutch 3481, GH, MO), Santa Elena on Fila Cotón, Cordillera de Talamanca (Davidse, Herrera & Grayum 28244 (MO), San Vito de Java (Raven 21776 (CR, F, GH, MO), Cerro Central de Zurquí (Dodge, J. Valerio R., Thomas & R. Valerio R. 6099, F, MO), La Hondura de San José (Valerio 751, F), Cordillera de Talamanca (Williams 19993 (F); especially on Atlantic slope in areas consistently bathed by clouds. Fl. Sept.—Jan. Guat—Pan., w. S.Am. to Peru and Bol. This is a very small, delicate epiphyte and, although common in Costa Rica, may be easily overlooked. Plants with flowers which have lost their corollas are frequently collected, yet can be readily identified by the small size of the plants and the conspicuous, broadly ovate sepals (chaffy in fruit) that are often as large or larger than the basal leaves.
Utricularia praetermissa P. Taylor, Ann. Missouri Bot. Gard. 63: 576. [1976] 1977.
Perennial, tubers present. Leaves 1-2(4) long-petiolate, 7-20 cm long, the lamina broadly to narrowly elliptic, sometimes broadly lanceolate to linear-lanceolate, mostly 3-8(10) cm long, 1-3 cm wide, coriaceous, tapering to a petiole. Traps numerous on capillary stolon branches (often broken off in collecting). Bracts straight, 5-8(10) mm long. Flowers 1-2(6), large. Corolla (2.5)3.5-5 cm long, white to pale pink or tinged with purple, yellow blotch at base of lower lip, glabrous or with a few marginal glandular hairs, lower lip nearly entire, rounded; spur curved upwards beneath lip and deflexed at apex (double curve), usually hidden beneath lower lip. Capsule ovoid, 8-9 mm long.
Epiphytic or terrestrial; on wet tree trunks and wet road banks in cloud forest areas. Mainly between 850 m and 2800 m elevation, sometimes lower. Region of San Ramón (Crow 7484, CR, F, MO, NHA), Monteverde Reserve (Haber ex Bello 3266, MO), region of Zarcero (Smith 498, F), Tapantí (Croat 36112, MO), Valle de Río San Bosco, Tilarán (Haber ex Bello, Clagget & Powell 5590, MO), Vara Blanca, Central Cordillera (Crow 7507, CR, F, MO, NHA), Volcán Barva (Burger & Baker 9457, CR, F), Monteverde Community, Pacific slope (Haber ex Bello 5298, MO), Osa Peninsula (Raven 21496, F), P. N. Braulio Carrillo (Crow & Rivera 7488, CR), Tinamaste (San Cristóbal) between San Isidro and Dominical (Maas & Cramer 1354, F), Fila División, Cuericí (Gómez 2352, CR, F),vicinity of El General (Skutch 2730, MO), Cascajal near Rios Cascajal and Bajo Máquina (J. & C. Taylor 11256, F). Fl. Jun.—Sept. Nic., C.R. and Pan. This is the most common and conspicuous epiphytic Utricularia in Costa Rica and is often mistaken for an orchid. It occurs primarily in open sites, mossy trees in pastures and forest margins and on steep, wet banks of road cuts with sparse vegetation (bladders usually located in bryophyte mats), where clouds bathe the area on a daily basis, but not in areas with strong, dessicating winds. According to Taylor (1989) notes that records of U. alpina from Panama northwards are based on misidentified specimens of U. praetermissa
Utricularia pusilla Vahl, Enum. 1:202. 1904.
Small annual, rhizoids present. Leaves numerous, 0.4-1 cm long, petiole tapering to narrowly obovate to spatulate lamina, 0.5-1(2) mm wide, 1-nerved, often decayed at anthesis. Inflorescence solitary, unbranched, filiform, 3-20 cm long. Bracts peltate, sterile bracts alternating with fertile, Flowers 2-15, small. Calyx lobes unequal, lower lobe slightly larger. Corolla 5-8 mm long, yellow, lower lip deeply 3-lobed, palate glandular and pubescent, swollen, bilobed; spur greatly exceeding the lower lip (often 2-3 times as long), decending. Capsule globose, 1.5-2 mm long.
Terrestrial; wet sandy sites in savannas. 200-1300 m elevation. Paraíso (Hernández & Sánchez 86017-2, CR), Turrialba (Oersted s.n., F), Lagunas Coyol y Guayabal, La Cruz (Gómez-Laurito 9169, CR), Río Liberia, near Hacienda Santamaría, (Dodge & Thomas 6261, MO), Sabana Dúrika (Weston 10218, F), Buenos Aires, Valle de El General (Gómez-Laurito 10127, CR, F), Agua Buena (Cañas Gordas) (Pittier 11084, CR), La Palma, Prov. San José (collector unknown, F). Fl. Jan., Feb., Sept. Mex.—Pan. S. to Argentina, Antilles. Utricularia pusilla is similar to the wide-ranging U. subulata, which has inflorescences lacking sterile bracts on the peduncle between flowers. Additionally the spur in U. pusilla is about twice as long as the lower lip, while in U. subulata the spur is about as long as the lower lip. It may also be confused with U. gibba when it is found flowering, stranded on mud (thus submersed portion not evident). However, U. pusilla is readily recognized by the peltate bracts, in contrast to the basifixed bracts of U. gibba.
Utricularia purpurea Walter, Fl. Carol. p. 64. 1788.
Perennial, submersed aquatic. Stolons long, slender, branched, all vegetative parts very mucilaginous. Leaves (modified branches) numerous, 4-whorled, repeatedly divided into whorled (or ultimately 2-3) capillary segments. Traps terminal on leaf segments. Inflorescence erect, emergent. Bracts peltate. Flowers 2-3(5) (one open at a time). Corolla 7-8(9) mm long (base to tip of lower lip), light purple to deep pink, yellow blotch at base of lower lip, lower lip 3-lobed, lateral lobes saccate; spur conical, apex obtuse, enclosed by inrolled lateral margins of lower lip. Capsule globose, ca. 3 mm diameter.
Submersed; shallow pools to deep lakes and slow moving waters. Low elevations. Known only from laguna at Buenos Aires, Valle de El General (Crow 6175, CR, F, MO, NHA; Grayum & Herrera 6066, MO). Fl. Sept.—Nov. E. Can. and U.S.A. S. to Florida and Tex., se. Mex., Belize, and CR, Cuba, Bahamas. This species is readily distinguished by its whorled leaves, thus it is not likely to be confused with the other purple-flowered species in Costa Rica, U. hydrocarpa.
Utricularia unifolia Ruiz & Pavon, Fl. Per. 1: 20. 1797.
Perennial, tubers present. Leaves 1-2(3) long-petiolate, up to 15 cm long, narrowly oblanceolate, the lamina narrowly elliptic, mostly 3-7 cm long, 1-2.5 cm wide, with very acute apex., coriaceous. Bracts long, curved, 10-30 mm long. Flowers 1-3, large. Corolla 3-4(5) cm long, lavender to light purple or violet, with white or yellow blotch at base of lower lip, glabrous or sparsely stipitate-glandular at the margin, lower lip nearly entire, rounded, palate with a pubescent, slightly raised marginal rim; spur subulate, apex curved upwards, shorter than lower lip. Capsule ovoid, 8-12 mm long.
Epiphytic or terrestrial; on wet mossy tree trunks and wet banks in cloud forest. Mainly 2000-3000 m elevation. Known only from Valle del Silencio, along Río Terbi, Cordillera de Talamanca, Atlantic Slope, near Panamá (Davidse, Herrera & Grayum 28586, CR, MO).