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Two new species of Burmeistera (Campanulaceae: Lobelioideae) from Antioquia, Colombia/Dos especies nuevas de Burmeistera (Campanulaceae: Lobelioideae) de Antioquia, Colombia.

INTRODUCTION

The neotropical Lobelioideae (Campanulaceae), especially the clade with the sister genera Burmeistera, Centropogon and Syphocampylus, have recently been recognized as exceptional in terms of rapid speciation rates that appear to be correlated to the uplift of the Andes, and the biotic agents involved in pollination and seed dispersal (Lagomarsino et al. 2016). With over 120 species distributed from Honduras to Peru, Burmeistera is one of the most diversified genera in this clade. The majority of species in the genus are terrestrial herbs to subshrubs, which often exhibit hemiepiphytic or climbing habits. Flower shapes and colors in Burmeistera are highly adapted primarily to bat pollination, but in some cases hummingbird or insect visitors have also been reported (Muchhala 2006, 2007a, b, Muchhala and Potts 2007). The most conspicuous synapomorphy for the genus is the dilated opening of the anther tube (Fig. 1). In contrast to the well known mechanisms of pollination in the genus, fruit or seed dispersal is less known (Gamba et al. 2017).

Until recently, the taxonomic studies of this genus, described by the Colombian botanist J. J. Triana (1854), were based upon herbarium specimens, but recent field expeditions to unexplored areas in Colombia have unveiled 11 new species described in a six year period (Garzon-Venegas and Gonzalez 2012, 2014, Garzon-Venegas et al. 2012, 2013, 2014). Two sets of collections primarily from the Parque Nacional Natural Las Orquideas, in the department of Antioquia (Colombia), have called our attention because they do not match any of the known species of Burmeistera from Colombia or neighboring countries (Wimmer 1943, 1953, 1968, McVaugh 1965, Wilbur 1975, 1976, 1981, Jeppesen 1981, Lammers 1998, 2002). Given that their traits are consistently found in the specimens at hand in the Herbario Nacional Colombiano, Universidad Nacional de Colombia, Bogota (COL), Herbario de la Universidad de Antioquia (HUA), Herbario de la Universidad Nacional de Colombia, sede Medellin (MEDEL), and the New York Botanical Garden herbarium (NY), we proceed to describe these two new species.

MATERIALS AND METHODS

The terminology used for the descriptions follows primarily Jackson (1900) and Hickey and King (2000). All the type specimens for the species similar to the new taxa here described were consulted through JSTOR (https://plants.jstor.org/) or directly examined at the herbaria COL, HUA, MEDEL and NY

RESULTS AND DISCUSSION

Burmeistera flstulosa F. Gonzalez & J. M. Velez, sp. nov. TYPE: Colombia. Antioquia: Urrao, corregimiento de Encarnacion, Parque Nacional Natural Las Orquideas, sector Calles, cerca de la cabana de Parques, margen izquierda del rio Calles, 6[grados]31'08" North, 76[grados]15'59" West, 1340 m, 7 abr 2011 (fl, fr), J. Betancur, J. M. Velez-Puerta, P. Pedraza, J. M. Orejuela & A. Duque 15048 (holotype MEDEL [MEDEL 56096], isotypes COL, MEDEL [56095], NY). (Figs. 1, 3)

Species similar to Burmeistera tomentosula H. Karst. & Triana, from which it differs by the glabrous surfaces, the leaves with petiole 1.4-2 cm long and lamina elliptic to ovate, glabrous beneath, the extremely reduced subtending leaves, the calyx lobes 2.5-3.5 x 0.8-1.2 mm, the corolla glabrous, the ventral anthers barbate, and the berries white and obconic to ellipsoid (versus pubescent surfaces, leaves with petiole 2-6 cm long and lamina elliptic to narrowly obovate, puberulous beneath, the subtending leaves not greatly reduced, the calyx lobes 5-10 x 2-4 mm, the corolla densely puberulous, the ventral anthers ebarbate, and the berries pink and subglobose in B. tomentosula).

Glabrous herbs to 1 m tall. Petiole 1.4-2 cm long, glabrous; lamina elliptic to ovate, (3.5-) 9.5-14 x (2.8-) 4-6 cm, base obtuse, round to slightly cordate, asymmetric, apex acute to shortly acuminate, margin finely serrulate, with 30-40 evenly distributed callose teeth on each side, glabrous on both surfaces, secondary veins 10-14 pairs, higher order venation conspicuously reticulate, submarginal vein present but discontinuous. Flowers densely arranged in a corymb in the distal portion of the branch with shortened internodes, each flower axillary to a glabrous, bracteiform, scaly or filiform reduced leaf to 4 x 1 mm. Floral peduncle 2-4.8 (6 in fruit) cm long, thick (in vivo), fistulous and pale yellow (in siccus), with a proximal and a distal constriction (likely functioning as abscission zones), ebracteolate. Hypanthium obconic, 4-5 x 2-2.5 mm, glabrous. Floral buds glabrous. Calyx lobes very narrowly ovate, 2.5-3.5 x 0.8-1.2 mm, forming broad (ca. 1 mm) sinuses between them, green; corolla 1.8-2.3 cm long, green, sometimes tinted maroon, glabrous, tube slightly inflated at its base, 8-12 mm long, ca. 1 mm in diam., lobes arranged at angles of 120-140[grados] with respect to the tube, slightly falcate, dorsal lobes 7-9 x 2.5-3 mm, lateral lobes 6.5-8 x 2.5-3 mm, ventral lobe 5.5-6.5 x 2-2.5 mm, inflated at its base; synandrium exserted by 5 mm from the sinus between the dorsal lobes; filament tube green, finely puberulous; anther tube green, ventral anthers barbate at preanthesis and early anthesis. Berries obconic to ellipsoid, 8-10 x 4-8.5 mm, glabrous, green when juvenile, then white, not inflated, crowned by the green, persistent and slightly acrescent (to 9 mm long) calyx lobes.

Paratypes. COLOMBIA. Antioquia: Urrao, corregimiento de Encarnacion, Parque Nacional Natural Las Orquideas, sector Calles, camino hacia La Virgen, 14 abr 2011 (fl, fr), J. Betancur, J. M. Velez-Puerta, P. Pedraza, J. M. Orejuela & A. Duque 15232 (COL, MEDEL); Frontino, corregimiento Nutibara, region of Murri, Alto de Cuevas, ca. 6[grados]45' North, 75[grados]20' West, 1500-1880 m, 16-18 oct 1987 (fl), J. L. Luteyn et al. 11699 (HUA, NY); Frontino, corregimiento Nutibara, region of Murri, Altos de Cuevas, ca. 6[grados]45' North, 75[grados]20' West, 1500-1880 m, 16-18 oct 1987 (fl, fr), J. L. Luteyn et al. 11717 (HUA, NY); Urrao, corregimiento de Encarnacion, Parque Nacional Natural Las Orquideas, vereda Calles, Parque Nacional Natural Las Orquideas, en el camino de la cabana a la quebrada El Agudelo, 6[grados]31'5" North, 76[grados]15'9" West, 1351 m, 26 ene 2011 (fl, fr), P. Pedraza-Penalosa et al. 1981 (COL); Urrao, corregimiento La Encarnacion, vereda Calles, Parque Nacional Natural Las Orquideas, finca de Alfonso Pino, divisoria de aguas entre las quebradas La Virola y El Bosque, al noroccidente de la cabana de Calles, 6[grados]31'35" North, 76[grados]15'50" West, 1450-1470 m, 27 ene 2011 (fl), P. Pedraza-Penalosa et al. 2030 (COL, NY); Frontino, Parque Nacional Natural Las Orquideas, vereda Venados Abajo, despues de la finca Vista Hermosa, divisoria de aguas entre el rio Venado y Quebrada La Selva, 6[grados]33'13" North, 76[grados]18'54.3" West, 1240-1280 m, 25 Jul 2011 (fl, fr), P. Pedraza-Penalosa et al. 2334 (COL).

Etymology. The specific epithet refers to the thick peduncle that becomes fistulose and pale yellow at late anthesis.

Distribution, habitat, conservation and phenology. Burmeistera fistulosa has been collected only in the department of Antioquia (Colombia). The species grows at the edges of primary or secondary forests between 1240 and 1880 m in elevation, including Quercus humboldtii forests. The species has been found setting flowers and fruits in January, April, July and October.

General comments. The new species is distinguishable by a corymbose flowering portion of the branchlets, and the fistulous floral peduncle with two well defined constrictions that likely facilitate fruit dispersal by providing break-joints allowing the fruit to be easily removed from the plant. These two traits resemble those found in Burmeistera tomentosula H. Karst. & Triana, from Cauca, Narino, Quindio and Valle del Cauca. However, they stand as two separate species, as they consistently differ in the following set of characters: Burmeistera fistulosa is glabrous, the petiole is 1.4-2 cm long, the leaf lamina is elliptic to ovate, glabrous beneath, the subtending leaves are extremely reduced to filiform to scaly bracts to 4 x 1 mm, the floral buds are glabrous, the calyx lobes are 2.5-3.5 x 0.8-1.2 mm, the corolla is glabrous, the ventral anthers are barbate, and the berries are white and obconic. In contrast, B. tomentosula is puberulous throughout, the petiole is longer (2-6 cm long), the leaf lamina is elliptic to narrowly obovate, puberulous beneath, the subtending bracts (1-2 x 0.4-0.8 cm) are not as reduced as in the former species, the floral buds are densely puberulous, the calyx lobes are 5-10 x 2-4 mm, the corolla is densely puberulous, the ventral anthers are ebarbate, and the berries are pink and subglobose.

According to Wimmer(1943), the corymbose arrangement of flowers is present in four species of Burmeistera, B. armadana E. Wimm., B. multiflora Zahlb., B. suerrensis E. Wimm. and B. virescens (Benth.) Hook. & B.D. Jacks. The first two species grow and are restricted to southern Colombia, whereas the latter two reach Ecuador. Jeppesen (1981) reduced B. armadana under the synonymy of B. multiflora. Besides its disjunct area of distribution, B. fistulosa differs by its ebracteolate, thick peduncle with a proximal and a distal abscission zone, and its obconic to ellipsoid, white berries, versus the bracteolate, slender peduncle that lacks abscission zones, and the globose, pink berries in B. multiflora.

Burmeistera reclinata F. Gonzalez & J. M. Velez, sp. nov. TYPE: Colombia. Antioquia: Frontino, Parque Nacional Natural Las Orquideas, vereda Venados Abajo, camino de la cabana de Venados a la vereda Alto Bonito, al frente de la cabana, 6[grados]32'10.6" North, 76[grados]19'15" West, 900-920 m, 24 Jul 2011 (fl, fr), P. Pedraza-Penalosa, J. Betancur, M. F. Gonzalez, R. Arevalo, D. Sanin, A. Zuluaga, A. Duque & J. Serna 2304 (holotype MEDEL, isotypes COL, NY) (Figs. 2, 3).

Species similar to Burmeistera diazii, from which it differs by the narrowly ovate leaves with 5-6 pairs of secondary veins and higher order veins inconspicuous beneath, the glabrous peduncle 3.3-6 cm long, the glabrous hypanthium 3-4 x 2-2.6 mm, the glabrous calyx lobes 5-9 x 0.6-1 mm, half of the corolla length or less, the glabrous corolla, the barbate ventral anthers at preanthesis, and the white berries suffused with vinaceous lines (versus ovate to lanceolate leaves with 7-8 (-12) pairs of secondary veins and higher order veins prominently reticulate beneath, the glabrous to sparsely puberulous peduncle (4.5-) 6.5-7.5 (-9) cm long, the glabrescent to puberulous hypanthium 5-7 x 5-6 mm, the glabrous to sparsely puberulous calyx lobes 13-16 x 2-3 mm, almost as long as the corolla tube, the sparsely puberulous corolla, the ebarbate ventral anthers at preanthesis, and the green berries tinted maroon in B. diazii).

Herbs to subshrubs with glabrous, reclinate, climbing or pendant branchlets to 2 m long. Internodes to 2.2 cm long, gradually reduced to 4 mm long between young leaves. Petiole 5-9 mm long, glabrous; leaf blade narrowly ovate, 5-6(-11) x 1.8-2.2(-2.8) cm, 3-4 times as long as wide, thick, glabrous on both surfaces, glaucescent on the lower surface, base obtuse to round, slightly asymmetric, apex acute to acuminate, margin subentire, with 5-12 callose, white (in vivo) intramarginal teeth evenly distributed on each side, main vein prominulous above, impressed beneath, second and higher order veins inconspicuous except by the 5-6 pairs of secondary veins slightly prominulous on the upper surface, submarginal vein lacking. Floral peduncle 3.3-6 cm long, 2-3 times longer than the leaves, glabrous, ebracteolate. Flowers 2-3 cm long. Hypanthium obconic, 3-4 x 2-2.6 mm; calyx lobes lineariform, 5-9 x 0.6-1 mm, half of the corolla length or less, patent to refiexed, leaving broad sinuses between them, margin with 2-3 teeth per side, green with vinaceous lines; corolla 2-2.6 cm long, 1-1.5 mm in diam., glabrous, uniformly green or yellowish green, or sometimes suffused with reddish lines, tube 1.1-1.3 cm long, slightly inflated at its base, lobes slightly falcate, arranged at angles of 140-160[grados] with respect to the tube, apex acute, dorsal lobes 9-11 x 1.5-2.5 mm, lateral lobes 8-9 x 1.5-2.3 mm, ventral lobe 8-8.5 x 1.5-2.2 mm, basally inflated; synandrium shortly exserted by 2-5 (-9) mm from the sinus between the dorsal lobes, puberulous; filament tube 1.5-2.3 cm long, green, anther tube purple, the ventral anthers with a dense tuft of long and curly white hairs during preanthesis that fall off during anthesis; stigma white. Berries oblate, 7-8 x 9-10 mm, spongy, not inflated, white suffused with vinaceous lines especially on the persistent and slightly accrescent calyx lobes that reach 10 x 2 mm, border of hypanthium yellow; seeds brown.

Paratypes. COLOMBIA. Antioquia: Urrao, corregimiento de Encarnacion, Parque Nacional Natural Las Orquideas, sector Calles, camino hacia La Virgen, 14 abr 2011 (fl), J. Betancur, J. M. Velez-Puerta et al. 15226 (COL, MEDEL); Frontino, road to Murri, 15.0 km W of Nutibara (Alto de Cuevas), margin of primary forest ca. 1 km S of road, 06[grados]45' North, 76[grados]23' West, 1850 m, 17 oct 1987 (fl, fr), A. E. Brant & G. E. Martinez 1345 (H|UA); Valdivia, vereda San Fermin, 1 km de la via Ventanas (Mun. Yarumal)-Briceno, fmca San Fermin, 1800-2000 m, 7[grados]15' North, 75[grados]30' West, 20 mar 1988 (fl, fr), R. Callejas et al. 6049 (HUA); Parque Nacional Natural Las Orquideas, sector Calles, margen derecha de la confluencia de los rios Polo y Calles, 1380 m, 6[grados]32' North, 76[grados]19' West, 26 mar 1988 (fl), A. Cogollo et al. 2627 (JAUM); Parque Nacional Natural Las Orquideas, sector Calles, margen derecha del rio Calles, 6[grados]32' North, 76[grados] 19' West, 1240 m, 29 mar 1988 (fl), A. Cogollo et al. 2794 (JAUM); Frontino, corregimiento Nutibara, Alto de Cuevas, quebrada El Oso, 1900 m, 22 abr 1991 (fl), R. Fonnegra et al. 3483 (HUA); to 5 km down road to San Fermin de Briceno, W off Pan American Hwy. ca. 25 km N of Yarumal, 1525-1830 m, ca. 7[grados]01' North, 75[grados]35' West, 26 may 1984 (fl, fr), J. L. Luteyn et al. 10774 (COL, JAUM, NY); Frontino, Nutibara-La Blanquita road, region of Murri, alto de rio Cuevas, ca. 9-18 km W of Nutibara, ca. 6[grados]45' North, 76[grados]20' Wes, 1450-2130 m, 10 apr 1989 (fl, fr), J. L. Luteyn et al. 12453 (COL, HUA, NY); Frontino, corregimiento Nutibara, Nutibara-La Blanquita road, region of Murri, alto de rio Cuevas, ca. 9-18 km W of Nutibara, ca. 6[grados]45' North, 76[grados]20' West, 1450-2130 m, 6-10 apr 1989 (fl, fr), J. L. Luteyn & S. Sylva 12471 (COL, HUA, NY); Frontino, corregimiento Nutibara, cuenca alta del rio Cuevas, 2020 m, 13 ene 1987 (fl), D. Sanchez et al. 919 (MEDEL, NY); Frontino, corregimiento Nutibara, cuenca alta del rio Cuevas, 1780 m, 23 sep 1987 (fl), D. Sanchez et al. 1612 (MEDEL, NY); km 1-5 on road to Briceno, c. 25 km N of Yarumal, 7[grados]07' North, 75[grados]28' West, 1850 m, 7 feb 1986 (fl, fr), B. A. Stein & A. Cogollo 3372 (COL, NY); Sonson, km 12 of road Sonson-Narino, 5[grados]42' North, 75[grados]15' West, 2630 m, J. L. Zarucchi & N. Bedoya 4428 (COL, HUA, NY); Frontino, km 13 of road Nutibara-La Blanquita, region of Murri, Alto de Cuevas, 06[grados]44' North, 76[grados]23' West, 1990 m, J. L. Zarucchi et al. 7223 (HUA, NY).

Etymology. The specific epithet refers to the reclinate, climbing or pendant branchlets.

Distribution, habitat, conservation status and phenology. Burmeistera reclinata has been collected in the Parque Nacional Natural Las Orquideas (Antioquia), and in the municipalities of Sonson and Valdivia. The species grows as a reclinate, climbing or pendant herb at cloud forest edges between 900 and 2630 m of altitude. It has been collected flowering and setting fruits from January through May, and in September and October.

General comments. Burmeistera reclinata is similar to B. diazii Garzon & F. Gonzalez as they both have narrow leaves that are glaucescent beneath (Table 1). At least eight additional species primarily from Central America also possess this trait; although vegetatively similar, the new species differs from the remaining species of this group by the unique combination of diagnostic traits (Table 1), mainly by the size of the flowers, and the relative size of the calyx lobes versus the corolla tube.

Some of the paratypes of Burmeistera reclinata were previously identified as B. crassifolia (E. Wimm.) E. Wimm., or as B. microphylla Donn. Sm., but a detailed comparison between the protologues and the type specimens of these species shows clear differences in both vegetative and reproductive traits (Table 1). These include the narrowly ovate leaves 3-4 times as long as wide, with inconspicuous venation and margin subentire to finely serrulate, the glabrous peduncle 3.3-6 cm, 2-3 times longer than the leaves (Fig. 2 b, c), the calyx lobes linear (5-9 x 0.6-1 mm), the corolla 2-2.6 cm long, with slightly falcate lobes, these arranged at angles of 140-160[grados] with respect to the tube, and the berries oblate, 7-8 x 9-10 mm (versus the elliptic, oblong, ovate or widely-ovate leaves that are 1-2 times as long as wide, the glabrous to puberulous peduncle 1.5-2 cm long, as long as the leaves, the corolla 1.26-2 cm long, the obovoid berries 1-1.2 x 1-1.2 cm in B. crassifolia; and the conspicuous leaf venation that is impressed on the upper surface and prominent on the lower surface, the crenate leaf margin, the oblong to ovate (c. 2 x 1 mm) calyx lobes, the corolla tube slightly inflated distally and the corolla lobes in an angle of ca. 90[grados] with respect to the tube in B. microphylla).

Burmeistera maculata E. Wimm. was described as a new species, based on F. W. Pennell 4440, from "Colombia: Dept. Bolivar, im Walde bei Antizales, 1700-2000 m", (Wimmer 1943:160), collected on February 25-26, 1918. The leaves of the holotype, deposited at NY are similar to those found in B. reclinata and the remaining species with glaucescent leaves (cf Table 1). Unfortunately, no isotypes are known, and the only flower in the specimen is severely fragmented to a small piece of the corolla; the calyx and the fruits, crucial to provide diagnostic and differential traits within Burmeistera, were not preserved. Thus, the identity of B. maculata is uncertain, as it could correspond to any of the species with glaucescent leaves, and it will remain as a species with "non satis notae", as it was originally published by Wimmer (1943).

AUTHOR'S CONTRIBUTIONS

Both authors equally contributed to the herbarium work and the writing of the manuscript. The second author also contributed to field observations and collections.

CONFLICT OF INTEREST

The authors declare no conflict of interest.

ACKNOWLEDGMENTS

The first author thanks the Faculty of Sciences, National University of Colombia, for granting a sabbatical year to write the monograph of Burmeistera for Flora of Colombia. Thanks to two anonymous reviewers for their valuable comments. We thank P. Pedraza-Penalosa (New York Botanical Garden), and J. Betancur, F Gomez and M. F. Gonzalez (National University of Colombia), for providing most of the photos that illustrate the two new species. We also thank N. L. Pabon-Mora (Universidad de Antioquia), and Sebastian Gonzalez (Universidad Nacional de Colombia) for preparing the figures here published. Finally, we thank the curators of COL, HUA, MEDEL and NY for continuously supporting this research.

LITERATURE CITED

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FAVIO GONZALEZ (1*), JORGE MARIO VELEZ-PUERTA (2)

(1) Universidad Nacional de Colombia (Sede Bogota). Facultad de Ciencias. Instituto de Ciencias Naturales. Cra. 30 #45-03, edificio 425, Bogota, Colombia. fagonzalezg@unal.edu.co

(2) Universidad Nacional de Colombia (Sede Medellin). Herbario MEDEl. AA 3840, Medellin, Colombia. jmvelezp@unal. edu. co

(*) Corresponding author.

Received: 16/02/2018

Accepted: 19/06/2018

https://dx.doi.org/10.15446/caldasia.v40n2.70457
Table 1. Comparison of diagnostic traits between Burmeistera reclinata
F. Gonzalez & J. M. Velez and nine closely related species with narrow
leaves, glaucescent beneath.

Trait           B. reclinata,    B. chiriquiensis
                sp. nov.         Wilbur (1976)

Branchlet       glabrous         glabrous
surface
Petiole         glabrous         glabrous
                narrowly
                ovate,
                5-6(-11)x
                1.8-2.2(-2.8)
                cm,              Very narrowly
                3-4 times        ovate, 10-12.5
Lamina          as long as       x 0.7-1 cm, ca.
shape           wide             10-15 times as
and size                         long as wide
                glabrous
                on both
                surfaces         glabrous on
                                 both surfaces
Lamina
indument        5-6 pairs of
                secondary
                veins, higher
                order veins
                inconspicuous    secondary veins
                                 inconspicuous
Leaf
venation
                3.3-6 cm,
                2-3 times
                longer than
                the leaves,
                glabrous
Peduncle                         ca. 3 cm, glabrous
length
and
surface         obconic, 3-4
                x 2-2.6 mm,
                glabrous
                2-3 cm
Hypanthium                       obconic, 5-6 x
                lineariform,     3 mm, glabrous
                5-9 x 0.6-1
                mm, half of
                the corolla      3-3.5 cm
Flower          length or
length          less, glabrous
                                 lineariform, 7-9
                                 x 0.5-0.8 mm,
                                 less than half
                                 of the corolla
                                 length glabrous
                2-2.6 cm,
Calyx           uniformly
lobes           green or
                yellowish
                green, or
                sometimes
                suffused
                with reddish     2.6-3 cm,
                lines, glabrous  greenish to
                                 dark purplish,
                11-13 mm,        glabrous
                slightly
Corolla         inflated at its
length,         base
shape,                           ca 14 mm,
color,          1.5-2.3 cm       inflated basally
and outer                        and distally
surface
                barbate
Corolla
tube            oblate, 7-8      ca. 2.3 cm
length          x 9-10 mm,
and             spongy, not
shape           inflated         barbate
Filament
tube
length                           fruits unknown
Ventral         Colombia
anthers         (Antioquia)
at preanthesis
Berries
                                   Panama (Bocas
                                   del Toro and
                                   Chiriqui)

                                                      B. diazii
                B. crassifolia                        Garzon &
Trait           (E. Wimm.) E.       B. darienensis    F. Gonzalez
                Wimm. (1943)        Wilbur (1976)     (2014)

Branchlet       glabrous            glabrous          glabrous
surface
Petiole         glabrous            glabrous          glabrous
                                                      ovate to
                                                      lanceolate,
                elliptic, oblong,   narrowly          (3-)4-6
                ovate or            elliptic to       (-7.5) x
                broadly ovate,      lanceolate,       (1.5-) 2.5-3
                3.5-10 x (1-)       (9-)12-17 x       (-4) cm, ca.
Lamina          3-5 cm, ca.         3-5 cm, ca.       2-3 times
shape           1-2 times as        3-4 times as      as long as
and size        long as wide        long as wide      wide
                                    glabrous          glabrous
                                    above,            on both
                glabrous on         puberulous on
                both surfaces                         surfaces
                                    the primary
                                    vein beneath      7-8 (-12)
Lamina                                                pairs of
indument                            12-16 pairs       secondary
                                    of secondary      veins, higher
                5-6 pairs ot        veins, higher     order veins
                secondary           order veins       prominently
                veins, higher       conspicuously     reticulate
                order veins         reticulate        beneath
Leaf            inconspicuous                         (4.5-) 6.5-7.5-9)
venation                                              cm,
                1.5-2 cm,                             glabrous
                as long as                            to sparsely
                the leaves,                           puberulous
                glabrous            3-5 cm, glabrous
                to sparsely
Peduncle        puberulous                            obconic, 5-7
length                                                x 5-6 mm,
and             obconic, 3-5                          glabrescent
surface         x 4-6 mm,                             to puberulous
                glabrous                              2.5-3 cm
                2-3 cm              cylindric, ca.    narrowly
                                    7 x 4 mm,         lanceolate,
Hypanthium                          glabrous          13-16 x
                lineariform,        ca. 4 cm          2-3 mm,
                4-10 x 0.5-1                          almost as
                mm, half of         ovate to          long as the
                the corolla         oblong, 2-3       corolla tube,
Flower          length or less,     x ca. 2 mm,       glabrous
length          glabrous            one third or      to sparsely
                                    less of the       puberulous
                                    corolla length
                                    glabrous
                                                      2-2.2
                                                      cm, green
                                                      suffused
Calyx           1.6-2 cm,                             with purple,
lobes           green suffused                        sparsely
                with purple,                          puberulous
                glabrous            1 2 - 1 5 cm
                                    dark red,
                                    glabrous          9-13 mm,
                                                      slightly
                11 - 16                               inflated
                inflated, at its                      basally
                base
                                                      ca. 2.3 cm
Corolla                             10 12 mm
length,         1.5-2.4 cm          inflated basally
shape,                              and distally      ebarbate
color,
and outer       barbate                               obconical,
surface                                               0.7-1 x
                obovoid, 1-1.2      ca. 2.5 cm        0.4-0.7 cm,
Corolla         x 1-1.2 cm,                           not inflated
tube            not inflated
length                              barbate
and
shape           Colombia                              Colombia
Filament        (Cauca,                               (Antioquia)
tube            Narino, valle       1.5 x 1.5 cm,
length          del Cauca)          not inflated
Ventral         and Ecuador
anthers         (Esmeraldas,
at preanthesis  Pichincha)
Berries                             ranama
                                    (Darien)

                 B. glauca
                 (E. Wimm.)      B. longifolia      B. tenuiflora
Trait            Gleason         Gleason            Donnell
                 (Wimmer         (1925)             Smith (1898)
                 1924)

Branchlet        glabrous        glabrous           glabrous to
surface                                             puberulous
                                 glabrescent
Petiole          glabrous        to puberulous      pubescent
                 lanceolate
                 to narrowly
                 ovate, 2.7-5.0                     elliptic to
                 (-8.0)                             oblong, 5-16
                 x 1.2-2.5                          x 2.5-5.5 cm,
                 cm, ca. 2-4                        mostly ca. 3
Lamina           times as        narrowly           times as long
shape            long as wide    ovate, 5-10        as wide
and size                         x 2-4 cm
                 glabrous
                 on both                            glabrous
                                                    above,
                 surfaces                           puberulous
                                 glabrescent
                                 above, puberulous  on the veins
Lamina           7-11 pairs      on                 beneath
indument         of secondary
                 veins, higher   the primary        7-9 pairs of
                 order veins     vein beneath       secondary
                 loosely                            veins, higher
                 reticulate      5-8 pairs of       order veins
                                 secondary          conspicuously
Leaf                             veins, higher      reticulate
venation                         order veins
                                 loosely
                                 reticulate
                 5-9 cm,
                 glabrous
                                                    4.5-8 cm,
Peduncle                                            glabrous
length
and                              5-7 cm,
surface                          glabrous
                 cylindric,
                 8-10 mm,                           cylindric
                 glabrous                           to obconic,
Hypanthium       3-3.5 cm                           5 x 3 mm,
                                 obconic,           glabrous
                 deltoid or      8-13 x 4-7         ca. 4 cm
                 ovate, 1.5-2.5  mm, sparsely
                 (-3.5)          puberulous
Flower           mm long,        3.8-4.2 cm
length           one tenth of    Narrowly
                 the corolla     oblong to          Lineariform,
                 length          narrowly           (7-) 11-20
                 glabrous,       ovate, 2-3.5       mm long,
                                 x 1-2 mm,          glabrous
                                 1/10 or less
                                 the length of
Calyx                            the corolla,
lobes            c. 2 cm,        puberulous
                 glabrous,       puberulous
                 greenish,
                 purplish or                        1.3-1.7 cm,
                 reddish                            glabrous
                                 to 3 cm,           or sparsely
                                 sparsely           pubescent
                 15-18 mm,       puberulous,        externally,
                 slightly        yellowish          greenish or
                 inflated        green              yellowish
Corolla          basally                            white
length,
shape,           ca. 2 cm        14-20 mm,          11-15 mm,
color,                           not inflated       inflated
and outer                                           basally and
surface          barbate                            distally
Corolla          globose, 3-6    ca. 3.7 cm         ca. 2.2 cm
tube             cm long x
length           (2-)3-5 cm,
and              inflated        ebarbate           ebarbate
shape
Filament                         subglobose,        obovoid,
tube                             2-3 x              7-10 x 5 mm,
length           Panama          1.5-2.6 cm,        inflated
Ventral          (Chiriqui)      inflated
anthers                                             Costa Rica
at preanthesis                                      (Cordillera de
                                                    Tilaran and
Berries                          Colombia           Cordillera
                                 (Antioquia,        Central), Panama
                                 Cordoba)           (Boquete
                                                    and Santa Fe
                                                    de Veraguas)

                B. toroensis
Trait           Wilbur (1976)       B. zurquiensis
                                    Wilbur (1975)

Branchlet                           glabrous
surface
                glabrous
Petiole                             sparsely ciliate

                glabrous
                                    lanceolate
                                    to narrowly
                                    ovate, 5-9.5 x
                                    (1.5-) 2.5-3.9
Lamina          elliptic, 4-7.5     cm, ca. 2-4
shape           x 1.5-2.7 cm,       times as long
and size        ca. 2.5-3 times     as wide
                as long as wide
                                    glabrous on
                                    both surfaces
Lamina
indument        glabrous on         6-8 pairs of
                both surfaces       secondary
                                    veins, higher
                                    order veins
                                    loosely reticulate
                5-7 pairs of
Leaf            secondary
venation        veins, higher
                order veins
                loosely reticulate
                                    2-5 cm, glabrous
Peduncle
length
and
surface
                ca. 5 cm,
                glabrous
                                    obconic, 5-8
Hypanthium                          x 4-7 mm
                                    glabrous
                                    4.5-5.5 cm
                obconic to
                cylindric, 6-8
Flower          x 4-5 mm,
length          glabrous
                3-4 cm              lanceolate,
                                    (6-)8-12 x 1.5-3
                                    mm, glabrous
Calyx
lobes           ovate, 6 mm,
                glabrous
                                    2.2-2.7 cm,
                                    glabrous,
                                    green, green
                                    suffused with
                                    maroon or
Corolla                             maroon
length,
shape,          1.5-1.7 cm
color,          glabrous,           20-25 mm,
and outer       purplish            slightly inflated
surface                             basally and
                                    distally
Corolla
tube                                4-4.5 cm
length          13-15 mm, not
and             inflated
shape                               barbate
Filament
tube                                globose, ca.
length          2-2.4 cm            9-11 x 9-11
Ventral                             mm, (not
anthers                             inflated?)
at preanthesis  barbate
Berries
                fruits unknown      Costa Rica
                                    (Heredia)
                Panama (Bocas
                del Toro/Chiriqui)
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Title Annotation:BOTANY
Author:Gonzalez, Favio; Puerta, Jorge Mario Velez
Publication:Caldasia
Date:Jul 1, 2018
Words:5115
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