Volume 10 NOVON Number 4 2000 A Synopsis of the Genus Blakea (Melastomataceae) in Mexico and Central America Frank Almeda Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-4599, U.S.A. [email protected] ABSTRACT. A regional synopsis of the genus Blak¬ students until recently. The genus has not been ea is presented that recognizes 33 species in Mex¬ treated in its entirety since Cogniauxs (1891) fa¬ ico and Central America. This summary includes a milial monograph. Even the most recent lloristic key, distributional and phonological information, treatments of the family Melastomataceae for the notes on morphological variation, and discussions Mesoamerican region appeared over 30 years ago that provide rationale lor relegating six binomials (Gleason, 1958; Standley, 1924, 1938; Standley & to synonymy. Descriptions and diagnoses are pre¬ Williams, 1963; Winkler, 1965) when many areas sented lor four new species of Blakea (B. colora- were still little-explored and poorly known botani- densis, B. darcyana, and B. perforata from Panama; cally. and B. usilsoniorum Irom southern Costa Rica and The account presented here is a precursor to the Panama). Analysis of the type specimens and re¬ treatment that will appear in Flora Mesoamericana. cently collected flowering material of Topobea stor- Four new species and one new generic transfer are kii necessitates its transler to Blakea. Three other proposed in the context ol a regional synopsis. This species (B. cuatrecasii, B. repens, and B. subcon- summary also includes a key to the 33 recognized nata var. obtusa), all ol which were previously species, distributional and phenological informa¬ known from Andean South America, are reported tion, notes on variation, and discussions that pro¬ from Panama for the first time. vide rationale for relegating previously recognized species to synonymy. Blakea, with approximately 100 species, is one of two berry-fruited genera comprising the natural Taxonomic Treatment neotropical tribe Blakeeae (Almeda, 1990). It is Blakea R Browne, Civ. Nat. Hist. Jamaica 323. best represented in wet montane regions from 1756. TYPE: Blakea trinervia P. Browne. southern Mexico and the West Indies to Bolivia and Brazil. Although northwestern South America is un¬ Trees, shrubs, or woody vines, often epiphytic doubtedly the center ol species richness for Blakea with glabrous or variously pubescent, terete or (Wurdack, 1973), southern Central America is quadrate branchlets. Leaves thick and coriaceous clearly the area ol local importance for unusual di¬ or chartaceous, 3- to 5-nerved or -plinerved with versity in floral morphology. This floral diversity re¬ the transverse secondary veins (when evident) flects adaptations to a broad array ol important pol¬ closely spaced, straight, and parallel. Flowers 6- linators in the region, including rodents, buzzing merous, axillary, solitary or fascicled, typically pe¬ bees, and hummingbirds (Almeda, 1990; Lumer, dunculate in the upper leaf axils and subtended by 1981; Lumer & Schoer, 1986). For a woody genus two pairs of decussate, free or variously fused, co¬ with showy, often spectacular, flowers, it is surpris¬ riaceous or foliaceous, sessile bracts inserted at the ing that Blakea has escaped the attention of serious base ol the hypanthium. Hypanthia campanulate or Novon 10: 299-319. 2000. 300 Novon hemispheric; calyx persistent or deciduous, rarely erally, bluntly obtuse or broadly rounded apically calyptrate, truncate or 2- to 6-lobate. Petals 6, with two typically well-separated (typically minute) white, pink, magenta, red, or some combination of pores. In Topobea the flowers also commonly have these colors, glabrous but often marginally ciliolate. 12 stamens, although several hexandrous and one Stamens 12, isomorphic and glabrous; anthers oval, octandrous species occur in Costa Rica and Pana¬ oblong, or elliptic, laterally compressed, bluntly ob¬ ma; the anthers of Topobea, in contrast to those of tuse, obconic, or broadly rounded at the summit, Blakea, are uniporose or biporose, linear-oblong to free or laterally connate with two typically well- oblong-subulate or rostrate, usually not compressed separated apical pores; connective thickened and laterally, and with approximate or confluent dorsal- unappendaged or dilated basally into a dorsal de- ly inclined broad pores. flexed, horizontal, or upturned spur. Stigma punc- The posture and orientation of stamens and the tiform to capitate. Ovary inferior, usually 6-locular. shape, length, and degree of fusion of floral bracts Fruit baccate; seeds ovoid, clavate, or narrowly pyr¬ are important diagnostic characters in Blakea. iform, testa smooth. Pressing and drying often conceal these features, frequently necessitating a dissection of hydrated Blakea and the closely related genus Topobea are material. This is especially important when analyz¬ the only two genera comprising the tribe Blakeeae. ing the species with large enveloping foliaceous flo¬ This tribe is characterized by axillary flowers that ral bracts. Because of the prevalence of epiphytism are individually subtended by two pairs of decus¬ in Blakea, coupled with the local distribution of sate bracts, baccate fruits, ovoid to pyramidal, many taxa, it seems likely that additional species smooth seeds, and wood with multiseriate rays and will be discovered as forest canopies and remote the frequent occurrence of druses (Almeda, 1990). areas are explored. Inadequate material of several Flowering material is essential for the unequivocal collections from Costa Rica and Panama probably generic placement of species in this tribe. In Blak¬ represent additional species for the area covered ea, each flower has 12 stamens; the anthers are by this study. Their identity will require evaluation biporose, oval, oblong, or elliptic, compressed lat¬ when better material is collected. Kky to the Species of Blakea in Mexico and Central America la. Flowers always pendent, petals imbricate or convolute when fully expanded to form a subcylindric tube-like corolla concealing the stamens, which form a ring around the straight style. 2a. Petals uniformly pink or magenta; floral bracts and calyx lobes red in color . 14. B. fuchsioides 2b. Petals green or green flushed with purple or reddish at the base; floral bracts and calyx lobes not red in color. 3a. Leaf blades modified adaxially at the petiole-laminar junction into flap-like saccate pouches, the basal and lateral margins of which are free from but conspicuously decurrent on the petiole. 4a. Pubescence of uppermost intemodes consisting of hairs 1—2 mm long; leaf blades bluntly denticulate; outer floral bracts linear-lanceolate, 15-21 mm long, equaling or exceeding the calyx lobes at anthesis; Costa Rica (Cerro Chompipe.Volcdn Barva, and Volcdn Irazu) . . . . 2. B. austin-smithii 4b. Pubescence of uppermost intemodes consisting of hairs mostly less than 0.5 mm long; leaf blades entire; outer floral bracts ovate to elliptic-ovate, 5.5—9 mm long, conspicuously short¬ er than the calyx lobes at anthesis; Costa Rica (Cordillera de Tilardn, Cordillera de Guan- acaste). 5. B. chlorantha 3b. Leaf blades not modified adaxially at the petiole-laminar junction into flap-like saccate pouches. 5a. Older petioles, floral peduncles, and bracts essentially glabrous, outer floral bracts 13—17 mm wide on fruiting hypanthia; Costa Rica (Cordillera de Talamanca and Volcdn Barva) . 23. B. penduliflora 5b. Older petioles, floral peduncles, and bracts moderately to copiously covered with oppressed or spreading brown hairs; outer floral bracts 5-9 mm wide on fruiting hypanthia. 6a. Uppermost intemodes and elevated primary nerves on abaxial leaf surfaces moderately to densely covered with spreading brown hairs less than 0.5 mm long; floral peduncles (2.0—)2.8—3.8 cm long; outer floral bracts 1.3—2.0 cm long; Mexico (Chiapas) and western Guatemala (Quezaltenango and San Marcos) . 25. B. purpusii 6b. Uppermost internodes and elevated primary nerves on abaxial leaf surfaces densely covered with appressed brown hairs 1-2.5 mm long; floral peduncles 1.2—2.0 mm long; outer floral bracts 0.7-0.8 cm long; western Panama (Chiriquf).17. B. gregii lb. Flowers erect or antrorsely divergent but never truly pendent, petals white, pink, magenta, red, or some combination of these colors, the petals well separated from one another or connivent when fully expanded but never forming a tube-like corolla that conceals the stamens. Volume 10, Number 4 Almeda 301 2000 Blakea in Mexico and Central America 7a. Calyx lobe apex elaborated into a deflexed ovate to cordate foliaceous appendage; style glandular- puberulent; cauline nodes lacking stipuliform setose flaps. 8a. Leaf blades (12.2—)19—35 X (10—)17—35.5 cm, cordate to ovate-orbicular; petioles 4.5—11.2 cm long; floral peduncles 0.7—1.6 cm long; anther sacs free from one another but connivent in a ring around the style. 31. B. tuberculata 8b. Leaf blades (6.3—)10.3—20.7 X 5.2—10.3 cm, elliptic to elliptic-ovate; petioles 1.5—3.7 cm long; floral peduncles 0.4—0.9 cm long; anther sacs laterally connate in a ring around the style .... .4. B. calycosa 7b. Calyx lobe apex not elaborated into a foliaceous appendage; style glabrous, or if glandular-p'uberulent, then uppermost cauline nodes bearing stipuliform setose flaps. 9a. Inner and outer floral bracts essentially free to the base. 10a. Adaxial leaf surface copiously covered with smooth or gland-tipped hairs; calyx lobes filiform . 32. B. wilburiana 10b. Adaxial leaf surface essentially glabrous; calyx lobes not filiform. 11a. Leaves at a node strongly dimorphic in size. 12a. Leaf blades subpeltate at the base, lacking inconspicuous domatia in the angles between the median vein and each of the two innermost veins on the abaxial surface. 18. B. guatemalensis 12b. Leaf blades not subpeltate at the base, inconspicuous domatia (these often rup¬ tured) typically formed in the angles between the median vein and each of the two innermost veins on the abaxial surface. 13. B. foliacea 1 lb. Leaves at a node not strongly dimorphic in size. 13a. Leaves sessile, auriculate to cordate-clasping at the base. 12. B. elliptica 13b. Leaves petiolate, not auriculate or cordate-clasping at the base. 14a. Outer floral bracts distinctly shorter than the hypanthium. 15a. Calyx fused and concealing petals in bud but rupturing at anthesis into 3^4 persistent or tardily deciduous lobes.LB. anomala 15b. Calyx lobes not fused and concealing petals in bud. 16a. Leaf blades bearing irregularly rupturing domatia in the angles between the median vein and each of the two proximal lateral veins on the abaxial surface; floral peduncles 1.3^4.8 cm long. 17a. Calyx lobes obsolete or evident only as depressed deltoid undulations; anther thecae laterally connate . .22. B. pauciftora 17b. Calyx lobes broadly ovate, 1-1.5 mm long and 2.5-3.5 mm wide basally; anther thecae free .... 19. B. hammelii 16b. Leaf blades lacking irregularly rupturing domatia in the angles between the median vein and each of the two proximal lateral veins on the abaxial surface; floral peduncles 0.5-1.2 cm long . 20. B. herrerae 14b. Outer floral bracts equaling or exceeding the hypanthium in length. 18a. Calyx consisting of a truncate flange, the lobes obsolete or merely low undulations. 19a. F loral peduncles 0.9—1.2 cm long; anther thecae free. . 1 1. B. darcyana, sp. nov. 19b. Floral peduncles 2.1 —4-.2 cm long; anther thecae laterally con¬ nate. 20a. Petals white with a flush of pink on the abaxial margins; anthers yellow, the apex rounded; hypanthium somewhat to markedly costulate in post-anthesis .15. B. gracilis 20b. Petals pink throughout; anthers pink or pale yellow-white, the apex bluntly obconic; hypanthium terete and smooth in post-anthesis. 21. B. litoralis 18b. Calyx consisting of well-defined lobes. 21a. Uppermost branchlet nodes bearing setose stipuliform flaps 1— 2 X 4—6 mm; style glandular-pubenilent. 26. B. repens 21b. Uppermost branchlet nodes lacking setose stipuliform flaps; style glabrous. 22a. Uppermost internodes quadrangular with carinate angles and interpetiolar ridges created by the basally vaginate petioles; leaf base acute to obtuse with the innermost pair of primary veins usually poculate-coalesced with the me¬ dian vein on the abaxial surface. . 29. B. subconnata var. obtusa 22b. Uppermost internodes essentially terete; leaf base obtuse to rounded with inconspicuous domatia (these often rup¬ tured) typically formed in the angles between the median 302 No von vein and each of the two innermost primary veins and sometimes in the angle of the outermost lateral veins as well on the abaxial surface. 13. B. foliacea 9b. Inner and/or outer floral bracts partially fused to completely fused for their entire length. 23a. Leaves sessile, auriculate to cordate-clasping at the base. 12. B. elliptica 23b. Leaves petiolate, neither auriculate nor cordate-clasping at the base. 24a. Nodes of upper branchlets swollen and often bearing small holes 1—2 mm diam. on the leaf scars that lead to domatia presumably occupied by ants . . 24. B. perforata, sp. nov. 24b. Nodes of upper branchlets not as above. 25a. Calyx fused and concealing petals in bud, the calyptra or lobes ultimately de¬ ciduous. 26a. Calyx calyptrate and falling away as a unit; the prominently lobed floral bracts superficially resembling the calyx; anthers 9.5—10 X 6-7 mm . . . . 7. B. costaricensis 26b. Calyx rupturing at anthesis into two to six ovate-oblong deciduous lobes, ultimately leaving a truncate flange or tube on fruiting hypanthia; anthers 5-7 X 2-4 mm. 27a. Flowers essentially sessile; petals 2-2.2 X 1.4—1.8 cm; anthers lat¬ erally connate, 5X2 mm.8. B. crinita 27b. Floral peduncles 0.9-3 cm long; petals 3.5-5.3 X 3^1 cm; anthers free, 7 X 3.5-4 mm .3. B. brunnea 25b. Calyx not fused and not concealing petals in bud, or if calyx is fused in bud, then the lobes rupturing regularly and persisting on fruiting hypanthia. 28a. Inner floral bracts free to the base; anther thecae laterally connate for at least half of their length. 9. B. cuatrecasii 28b. I nner floral bracts iused basally for at least hall of their length; anther thecae free. 29a. Petals brilliant red with a small white band at the base; anther con¬ nective prolonged dorsally into an upturned spur 3-5 mm long . . . .27. B, scarlatina 29b. Petals white, pink, or some combination of these colors; anther con¬ nective not prolonged dorsally into an acute spur. 30a. Inner floral bracts connate for essentially their entire length to form a cupulate truncate collar that closely envelops the hy- panthium. 31a. Outer floral bracts fused for hall or more of their length and closely enveloping inner floral bracts and the hypan- thium; fruiting hypanthia 1.5-2 cm long. 32a. Outer floral bracts exceeding the inner floral bracts in length and typically concealing them on flowering and fruiting hypanthia; peduncles on fruiting hy¬ panthia 1.2-2 cm long . . . 33. B. wilsoniorurn, sp. nov. 32b. Outer floral bracts somewhat shorter than the inner floral bracts, the distal portion of inner bracts visible on flowering and fruiting hypanthia; peduncles on fruiting hypanthia 2.5-5 cm long. 28. B. storkii 31b. Outer floral bracts fused for less than half of their length, the lobes spreading and not closely enveloping inner floral bracts or the hypanthium; fruiting hypanthium 0.5—0.7 cm long. 6. B. coloradensis, sp. nov. 30b. Inner floral bracts connate for only a portion of their length to form a 2-lobed collar that only partially envelops the hypan¬ thium. 33a. Floral peduncles (1.5—)2.7—4 cm long; petals 2.8-3.8 X 2-3.2 cm, white only flushed with pink apically and along the margins. 16. B. grandiflora 33b. Floral peduncles 0.4—1.2 cm long; petals 1.5—2.5 X 0.7- 1.8 cm, entirely pink or white. 34a. Uppermost nodes beset with an interpetiolar tuft of coarse hairs that falls away to leave a distinct in¬ terpetiolar ridge or line; leaf blades not white-punc- ticulate adaxially when dry; outer floral bracts fused basally for 3-6 mm; petals white; Costa Rica .... . 30. B. taparuiana 34b. Uppermost nodes not beset with an interpetiolar tuft of coarse hairs that falls away to leave a distinct Volume 10, Number 4 Almeda 303 2000 Blakea in Mexico and Central America interpetiolar ridge or line; leaf blades white punc- ticulate adaxially when dry; outer floral bracts fused basally for 7—10 mm; petals pink; Belize, Guate¬ mala, and Honduras . 10. B. cuneata 1. Blakea anomala Donnell Smith, Bot. Gaz. 42: Distribution and phenology. Endemic to Costa 297. 1906. TYPE: Costa Rica. San Jose: In Rica where it is largely restricted to cloud forests truncis putridis ad La Palma, 1450-1550 m, on slopes of the Cordillera Central at 1500-2400 Sep. 1896, Pittier 10165 (syntype, US; isosyn- m. Collected in flower from November through types, BR, CR, NY); Costa Rica. San Jose: In March, in fruit from February through April. truncis putridis ad La Palma, 1450-1550 m, Among the species of Blakea with pendent green Sep. 1896, Tonduz 12521 (syntype, US; iso- flowers, B. austin-smithii is distinguished by its syntype, NY). saccate pouches at the petiole-laminar junction, Distribution and phenology. Endemic to Costa bluntly denticulate leaf blades, and linear-lanceo¬ Rica where it can be locally common in cloud for¬ late outer floral bracts that equal or exceed the ca¬ lyx lobes at anthesis. ests ol the Cordillera Central and Cordillera de Tal- amanca at 700—1800(—2400) m. Collected in flower from September through April, fruiting into August. 3. Blakea brunnea Cleason, in Woodson & Blakea anomala is readily recognized by its ir¬ Schery, Ann. Missouri Bot. Gard. 28: 435. regularly rupturing calyx, small scalelike floral 1941. TYPE: Panama. Chiriquf: vicinity of bracts, and laterally connate anthers. This species Bajo Chorro, 1900 m, 20-22 July 1940, Wood- exhibits two modal tendencies with respect to pu¬ son & Schery 618 (holotype, NY; isotype, MO). bescence and shape of the leaf base. Tbe syntypes Distribution and phenology. Endemic to the (Pittier 10165 and Tonduz 12521) have copiously cloud forests of western Panama at 900-2300 m. setose-furfuraceous upper internodes with leaves Flowering collections have been made in January, that also have varyingly persistent pubescence March, and August, fruiting collections in July and abaxially, and leal bases that are subcordate to December. rounded. This form varies through a series of less The flowers of this species, which measure 13.5 pubescent intermediates to individuals that are cm across when fresh, are the largest of any known nearly glabrous or at most scurfy puberulent on member of the Melastomataceae in the Mesoamer- vegetative buds with leaves that are mostly basally ican region. The large flowers, rupturing calyx, and acute. The glabrous populations also have conspic¬ mixture of stout conic and flattened hairs on floral uous elevated interpetiolar ridges on the uppermost peduncles and young vegetative organs readily dis¬ twigs, but these are lacking or not readily evident tinguish this species. in the pubescent populations represented by the syntype material cited herein. Judging from avail¬ 4. Blakea calycosa Gleason, Phytologia 1: 342. able collections examined for this study, the gla¬ 1939. TYPE: Panama. Code: vicinity of El brous variant is the more widespread form. The dif¬ Valle, 600-1000 m, 14 May 1939, Allen 1788 ferences exhibited by these two extremes do not (holotype, NY; isotypes, F, MO, US). seem to correlate with other characters, nor do they appear to be correlated with geographic or eleva- Distribution and phenology. Local and uncom¬ tional differences. The extremes when compared mon in cloud forests of Costa Rica and Panama at side by side are impressive, but they are bridged 600—1400 m. Flowering and fruiting collections by many intermediates that make formal taxonomic have been made from August through March. recognition pointless and arbitrary. Therefore, the Blakea calycosa is most similar to B. tuberculata species as recognized here is considered variable with which it shares several diagnostic characters and polymorphic. Perhaps the observed differences such as tuberculate petals, glandular-pul>erulent are the result of repeated hybridization between two styles, reflexed foliaceous appendages at the tip of formerly distinct entities. each calyx lobe, and similar anther appendages. They share a similar indument on young vegetative 2. Blakea austin-smithii Standley, Field Mus. buds that consists of variously roughened subulate Nat. Hist., Bot. Ser. 18: 1561. 1938. TYPE: hairs. After examining types and numerous collec¬ Costa Rica. Alajuela: Palmira, 2200 m. Mar. tions of both taxa, I have concluded that these two 1938, A. Smith H382 (holotype, F; isotype, species are closely related but distinct and worthy MO). of continued recognition. Diagnostic character dif- 304 Novon ferences include foliar shape and size (shorter and bra, 5-nervata, nervis secundariis 1.5—2 mm inter se distantibus. Flores 6-rneri in quoque nodo superiore 1-2, narrower in B. calycosa), petiole and petal length peduncularis 2^1.5 cm longis; bracteae exteriores 4—6 X (longer in B. tuberculata), free vs. laterally connate 9—10 mm ovatae ad basim paulo (1.5—2 mm) coalitae; anther sacs (in B. calycosa), and staminal append¬ bracteae interiores 6-7 mm retusae vel subemarginatae age posture (deflexed in B. calycosa vs. upturned omnino coalitae. Hypanthium (ad torum) 5—7 mm longum; in B. tuberculata). The taxonomy of this species calycis tubus ca. 3 mm longus, lobis 3^4 X 4—6 mm ro- tundatis usque truncatis. Petala 1.7—1.9 X 1.6-1.8 cm pair is complicated, however, by a series of some obovata apice rotundato. Antherae 4—5 X 1-1.5 mm ob- 25 anomalous collections from Costa Rica and Pan¬ longae inter se lateraliter non eohaerenles apice minute ama that do not fit comfortably into either taxon. A biporosae; connectivum ad basim dorsaliter paulo eleva- collection from the Caribbean slope of Costa Rica, tum. Ovarium 6-loculare, cono glabro (collo non evoluto). for example (de Nevers et al. 7797, CAS), has leaf Epiphytic shrub 2-3 m tall. Upper branchlets blades that are shaped like B. calycosa but ap¬ glabrous and sparingly lenticellate, distinctly quad¬ proach B. tuberculata in overall size. In flower size rate to sulcate becoming rounded-quadrate with this collection is intermediate between the two taxa, age. Vegetative buds, young petioles and floral and it has the prominent ovary cone anti collar of buds, and the adaxial surface and elevated primary the latter. Another collection (McPherson 7183, veins of immature leaves moderately to densely CAS) from Cerro Tute, Panama, has petioles that covered with a brownish scurfy indument of mealy approach B. tuberculata in length, leaf blades that ill-defined matted particles. Mature leaves of a pair are intermediate in size and shape, flowers that are equal to somewhat unequal in size, essentially gla¬ intermediate in size, and an ovary apex that is mod¬ brous on both surfaces; petioles 0.6-1.7 cm long; ified into a low elevated cone surrounded by widely blades coriaceous, 6.4-12.5 X 3.2-5.8 cm, elliptic, spaced glandular hairs reminiscent of B. calycosa. apex acuminate, base acute varying to obtuse, mar¬ Field studies are needed to better evaluate the sta¬ gin entire to subentire, 5-nerved, the transverse tus of these enigmatic populations and to determine secondary veins spaced 1.5—2 mm apart at the wid¬ whether hybridization and introgression are respon¬ sible for the observed patterns of variation. est portion of the blade. Flowers erect, 1 or 2 in each leaf axil of uppermost branches; peduncles 2— 4.5 cm long, rounded-quadrate, glabrous and len¬ 5. Blakea chlorantha Almeda, Rhodora 82: 609. ticellate. Floral bracts sessile, outer bracts 4—6 X 1980. TYPE: Costa Rica. Puntarenas: 3—3.5 9-10 mm, fused basally for 1.5-2 mm, flaring to km SE of Santa Elena and 2—3 km E of Mon- form a skirtlike collar, depressed-ovate, apex blunt¬ teverde on the Pefias Blancas trail. Cordillera ly retuse to subemarginate, margin entire but ± de Tilar^n, 1500-1540 m, 19 Dec. 1973, Al¬ undulate-involute, 3-veined, glabrous throughout; meda et al. 2005 (holotype, DUKE; isotypes, inner bracts connate for their entire length to form CAS, CR, MO, NY, US). an unlobed cuplike collar 6—7 mm long that closely Distribution and phenology. Endemic to Costa envelops the hypanthium. Hypanthium (in post an¬ Rica where it is confined to the Cordillera de Ti- thesis) eampanulate, 5-7 mm long to the torus and lardn and adjacent areas in cloud forest, elfin wood¬ 8—10 mm diam., glabrous. Calyx tube ca. 3 mm land, and windswept ridges at 900-1800 m. Col¬ long; free portions of calyx lobes 3-4 mm long and lected in flower from October through May, in fruit 4—6 mm wide basally, rounded-truncate, margin from January through May. coarsely entire and somewhat callose-thickened at This species is readily recognized by its semi¬ the median apex, glabrous on both surfaces. Petals circular pseudoformicarial pouches at the petiole- 6, glabrous, 1.7—1.9 X 1.6—1.8 cm, white, obovate, laminar junction, entire leaf blades, and ovate to apically rounded, entire. Stamens 12, isomorphic; elliptic-ovate outer floral bracts that are conspicu¬ filaments 6 mm long, complanate, glabrous and ously shorter than the calyx lobes at anthesis. somewhat declined to one side of the flower oppos¬ ing the style; anthers free, 4—5 mm long, 1-1.5 mm 6. Blakea coloradensis Almeda, sp. nov. TYPE: wide, yellow, oblong, laterally compressed with two Panama. Chiriquf/Bocas del Toro border: cloud well-separated pores at the rounded-truncate apex; forest off road to Cerro Colorado, 1450 m, 26 connective dilated dorso-basally ca. 0.5 mm above Jan. 1989, Almeda et al. 6417 (holotype, CAS; the filament insertion into a blunt appendage. Ova¬ isotypes, MEXU, MO, NY, PMA, US). Figure 1. ry 6-celled, glabrous at the slightly elevated bluntly conic apex but lacking a collar. Style erect and Frutex epiphyticus 2—3 m altus. Petioli 0.6-1.7 cm lon- gi; lamina 6.4—12.5 X 3.2—5.8 cm elliptica apice acumi¬ somewhat incurved distally, glabrous, 1.1-1.3 cm nata basi acuta vel obtusa ad maturitatem coriacea et gla¬ long, the expanded stigma subtruncate to broadly Volume 10, Number 4 Almeda 305 2000 Blakea in Mexico and Central America Figure 1. Blakea coloradensis Almeda. —A. Habit. —B. Representative leaf (abaxial surface). —C. Open flower with several petals and outer bracts removed. —I). Hypanthium and floral bracts with petals, androecium, and style removed. E. Petal (adaxial surface). —E Stamens, ventral view (left), lateral view (right). —G. Hypanthium (top view) showing ovary summit, torus, and calyx lobes. (A-G from Almeda et al. 6417.) 306 Novon rounded. Berry subglobose, 1 X 1 cm. Seeds beige, ering collections have been made in April; fruiting mostly 1 mm long. collections are unknown. Among Mesoamerican species of Blakea, this Distribution and phenology. A little-collected species is unusual in having large flowers that mea¬ cloud forest species presently known from the Pa¬ sure 10-11 cm across when fresh, a calyptriform cific slope of Cerro Colorado, Panama, at 1390— calyx, and floral bracts that are 6-lobed and remi¬ 1450 m. Collected in flower in January, in fruit in niscent of a typical calyx. Correa & Montenegro July. 10169 and McPherson 7926 (both at CAS) from Blakea coloradensis is unique among its Me- Parque Nacional Altos de Campana, Panama, are soamerican congeners in having outer floral bracts reminiscent of B. costaricensis in having thick co¬ that are widely spreading to form a saucer-like riaceous leaves and a calyptriform calyx. Field ob¬ structure at the base of the hypanthium. The pro¬ servations and additional study of the Panama nounced spreading posture of the outer floral bracts plants are needed to determine whether they are exposes the completely fused inner floral bracts in conspecific with B. costaricensis. a way that is never seen in other members of the genus with well-developed foliaceous outer bracts. 8. Blakea crinita Gleason, Phytologia 3: 356. This distinctive bracteal feature coupled with the 1950. TYPE: Panama. Code: crest of Cerro short hypanthium (5—7 mm long), comparatively Pajita, Allen 3956 (holotype, NY; isotypes, narrow leaves (3.2-5.8 cm wide), and small petals BM, MO). (1.7-1.9 X 1.6-1.8 cm) readily sets B. coloradensis apart from all other members of the genus. Distribution and phenology. This little-collect¬ In having dorsally appendiculate anthers and in¬ ed species is known only from cloud forest habitats ner floral bracts that are completely fused to form on Cerros Pajita and Gaital in the vicinity of El an apically truncate, cupulate collar that tightly en¬ Valle de Anton, Panama, at 900-1400 m. Collected velops the hypanthium, B. coloradensis is most sim¬ in flower from November through January, in very ilar to B. storkii of Costa Rica and Panama. Plants young fruit in February, July, and November. of the latter species are invariably coarser with out¬ In the protologue of this species, Gleason stated er floral bracts that are fused for half or more of that he could not satisfactorily determine the struc¬ their length to form a cup that closely envelops the ture of the hypanthium and involucre (floral bracts) inner bracts and hypanthium, the hypanthium because of the dense hair covering. This species proper is longer (1.2-1.9 cm), and the petals are was known to him only from the type. Two of the modally larger (2-4.2 X 1.5—2.9 cm). Since B. co¬ five additional collections examined for this treat¬ loradensis appears to be a local endemic with a ment each have a single open flower and none has limited elevational range, it is tempting to suggest mature fruits, so a thorough analysis of the repro¬ that it was derived from an allopatric relative like ductive characters in this species is still needed. B. storkii, which occurs at mostly higher elevations Blakea crinita is unusual in having a combination (1500—2636 m) from Cerro Pate Macho in western of essentially sessile flowers, laterally connate an¬ Panama northwestward to the Cordillera Central of thers, and a calyx that ruptures at anthesis in two Costa Rica. to six deciduous lobes. On fruiting hypanthia, rem¬ nants of the calyx form a tubelike flange. Paratype. PANAMA. Chiriqiif: Cerro Colorado, along road to copper mine, 34.1 km beyond bridge over Rio San 9. Blakea cuatrecasii Gleason, Bull. Torrey Bot. Felix near town of San Felix, 13.1 km beyond turnoff to Club 72: 387. 1945. TYPE: Colombia. Valle: Escopeta, 15 July 1976 (fr), Croat 37300 (MO). Cordillera Occidental, 900—1180 m, Cuatre- casas 15180 (holotype, NY; isotypes, CAS, F, 7. Blakea costaricensis Umana & Almeda, Sel- US). byana 12: 1. 1991. TYPE: Costa Rica. Limon: Distribution and phenology. Localized in cen¬ Valle de Estrella, Fila Matama, El Progreso, tral and eastern Panama where it is known from 1600 m, 24 Apr. 1989, Herrera & Chacon cloud forests on Cerros Jefe, Mali, Sapo, and Pirre 2758 (holotype, CR; isotypes, CAS, COL, F, extending to western Colombia at 650-1500 m. K, MEXU, MO, QCNE, US, USJ, WIS). Collected in flower and fruit sporadically through¬ Distribution and phenology. This little-collect¬ out the year. ed endemic of Costa Rica appears to be restricted This species exhibits much variation in leaf to the Caribbean slopes of the Cordillera de Tala- shape and the size and degree of fusion of the outer manca in montane forests at 1300—1600 m. Flow¬ floral bracts, but none of the differences can be Volume 10, Number 4 Almeda 307 2000 Blakea in Mexico and Central America correlated with geography or other characters. The apice minute biporosae; connectivum ad basim dorsaliter recognition of a single taxonomic entity seems war¬ dente 0.25 mm descendenti armatum. Ovarium 6-loculare, cono glabro (eollo non evoluto). ranted in view of interpopulational homogeneity with respect to the lepidote indument on abaxial Shrub. Upper branchlets terete to rounded-quad¬ foliar surfaces, semicircular floral bracts, bicolored rate and glabrous with thickened interpetiolar lines petals (white with red-pink apical band abaxially), or ridges. Mature leaves of a pair equal or nearly and pseudoterminal position of the anther append¬ so, glabrous on both surfaces; petioles 1.1-2.1 cm ages. long; blades coriaceous when dry, 4.9-7.7 cm long and 3-4.5 cm wide, obovate, apex caudate, base 10. Blakea cuneata Standley, Publ. Carnegie acute to acuminate, margin entire, 3- to 5-nerved Inst. Wash. 461: 76. 1936. TYPE: Belize. Pe- with inconspicuous but well-developed, irregularly ten: Rio Viejo, Camp 32, 20 Nov. 1933, Schipp ruptured domatia formed at the abaxial blade base S-604 (holotype, F). in the angles between the median vein and each of the two proximal lateral veins, the striolate trans¬ Blakea bella Standley, Field Mus. Nat. Hist., Bot. Ser. 22: 94. 1940. Syn. nov. TYPE: Guatemala. Izabal: hank verse secondary veins spaced 0.25 mm apart or less of Rio Du Ice, 28 Mar. 1939, Wilson 402 (holotype, at the widest portion of the blade. Flowers erect, F). solitary in the uppermost leaf axils and appearing Distribution and phenology. Local in rainforests pseudoterminal; peduncles 0.9—1.2 cm long, some¬ from Belize to Guatemala and Honduras at 250- what compressed laterally when dry, glabrous 1140 m. Collected in flower during September and throughout. Floral bracts sessile, entire and gla¬ October, in fruit during April and May. brous throughout; outer bracts 1.1-1.3 X 1.1-1.2 The only character used to distinguish B. cunea¬ cm, free or fused basally for 1 mm at anthesis, ovate ta and B. bella in the past has been petal length to elliptic-ovate, apex bluntly short-acuminate; in¬ (Standley & Williams, 1963). Petal size for the for¬ ner bracts 1.1—1.2 X 1—1.4 cm, free, obovate, apex mer was reported as 1.5 cm long in the protologue, rounded. Hypanthium campanulate to suburceola- whereas that of the latter was given as 3 cm long. te, 0.6 cm long to the torus and 0.6 cm diam., gla¬ Petals on the holotype of B. bella are in fact only brous. Calyx 4 mm long from the torus, flangelike 2-2.5 cm long. All other specimens attributable to and involute, the lobes merely depressed undula¬ this complex have petals ranging in length from 1.5 tions with inconspicuous callose thickenings at the to 2.5 cm. Because these two species are otherwise median abaxial apex, glabrous on both surfaces. identical or exhibit comparable patterns of size var¬ Petals 6, glabrous, 12-13 X 13.5-17 mm, white iation in all other features, there appears to be no flushed with pink, depressed-ovate to suborbicular, sound basis for the continued recognition of two apically rounded to gently undulate, entire. Sta¬ taxa. I herein relegate B. bella to the synonymy of mens 12, isomorphic, filaments 5-8 mm long, com- B. cuneata. planate and glabrous; anthers free, 4.5-5 mm long, 3-3.5 mm wide on the lateral face, probably yellow, ellipsoid in profile view, laterally compressed with 11. Blakea darcyana Almeda, sp. nov. TYPE: two well-separated pores at the apex; connective Panama. Chiriqui: Fortuna Dam area, N of res¬ thickened and dilated dorso-basally into a blunt de- ervoir, ridge along Continental Divide and flexed appendage less than 0.25 mm long about 1 southward from Quebrada de Arena, 1100- mm above the base of the anther thecae. Ovary 6- 1500 m, Aug. 1984, W. G. D'Arcy & C. Todzia celled, glabrous at the elevated conelike apex. Style 15958 (holotype, CAS; isotypes, MO, US). Fig¬ ure 2. glabrous, 6—7 mm long; stigma punctiform. Mature berry and seeds not seen. Frutex. Petioli 1.1—2.1 cm longi; lamina 4.9—7.7 cm obovata apice caudata basi acuta vel acuminata ad ma- Distribution and phenology. Known only from turitatem eoriacea et glabra, 3-5-nervata, nervi in axillis the type collection made in western Panama. acarodomatiis instructi, nervis secundariis 0.25 mm inter Blakea darcyana is distinguished by its glabrous, se distantibus. Flores 6-meri in quoque nodo superiore singuli, peduncularis 0.9-1.2 cm longis; bracteae exter- apically caudate leaves, finely striolate foliar ve¬ iores 1.1—1.3 X 1.1—1.2 cm ovatae vel ovato-ellipticae, ad nation, foliaceous floral bracts that conceal the hv- basim paulo (1 mm) coalitae vel omnino liberae; bracteae panthium, flangelike involute calyx tube, and dor¬ interiores 1—1.2 X 1 — 1.4 cm obovatae, omnino liberae. so-basally appendiculate anther connective. Hypanthium (ad torum) 0.6 cm longum; calycis tubus 4 Another distinctive, but readily overlooked, diag¬ mm longus, lobis paullulo 6-undulatus. Petala 12-13 X 13.5- 17 mm ovata apice rotundato vel undulato. Antherae nostic feature of this species is the domatia (most 4.5— 5 X 3—3.5 mm inter se lateral iter non cohaerentes evident when ruptured) at the abaxial blade base 308 Novon Figure 2. Blakea darcyana Almeda. —A. Habit. —B. Representative leaf (abaxial surface). —C. Leaf base (abaxial surface) showing perforated domatia. —D. Flower bud enveloped and concealed by outer and inner bracts. —E. Flower in pre-anthesis with three petals removed. —F. Petal (adaxial surface). —G. Stamen, lateral view. (A-G from D’Arcy & Todzia 15958.)