ON THE TYPIFICATION OF BURMEISTERA ( CAMPANULACEAE-LOBELIOIDEAE ) AND THE IDENTITY OF B

The genus Burmeistera, described by J. J. Triana in 1854, was typified with the species B. ibaguensis. The type specimen of this species consists of three fragments, two of which correspond to the type species and the third fragment belongs to B. rivina. Here we clarify the typification of the genus and the two species mentioned, and revise the descriptions of these species. In addition, a lectotype is here designated for B. rivina, and the name B. ceratocarpa var. dentata is subordinated to synonymy.


INTRODUCTION
With approximately 110 species (Lammers 2007, Garzón & González 2014), Burmeistera Triana is the fourth largest genus of the subfamily Lobelioideae.Their species can be distinguished from those of their related genera Centropogon C. Presl.and Siphocampylus Pohl by the broad (versus narrow) apex of the anther tube, the fleshy, indehiscent fruits (versus capsules in most Siphocampylus spp.), and the fusiform (versus ellipsoid to oblate) seeds.The genus was described by the Colombian botanist José Jerónimo Triana (1854), based on the species B. ibaguensis.Until now, the location of the type specimen and the identity of B. ibaguensis were in doubt, as it was stated in the following Editor's note by Tracy Elliot Hazen in Gleason's (1925:102) synopsis of the genus: "In view of Dr. Gleason's characterization of G. (sic) lacerata as 'so common a plant,' it is a matter of surprise and disappointment to the editor, as a member of the Colombian Expedition of 1922, that this species was not found by us in the type region, the Quindio, where much time was spent in intensive collecting.It is even more disappointing that though two members of the Expedition, in going over the Old Quindio Trail to Ibagué, passed through «La Pamilla,» (sic) the type station for Karsten›s unknown B. ibaguensis, no Burmeistera was found on that trip.This failure can hardly be ascribed entirely to the fact that this genus is characterized by inconspicuous green flowers".
The taxonomic uncertainty of Burmeistera ibaguensis was also recognized by Wimmer (1943:144), who included this species as one of the three poorly known species of Burmeistera ("Burmeisterae non satis notae"), all of which appear to be known to him only from the original descriptions.
During the ongoing taxonomic revision of Burmeistera for the Flora of Colombia, we have located the type specimen of B. ibaguensis, collected by Triana under the number 3069.1.1/88,and deposited in BM (Fig. 1).The label indicates that the specimen was collected in the "Prov.de Mariquita, La Palmilla, Alt.2300 m, Febr.1853," a locality that coincides the original description ("lugares sombríos del Monte del Quindío, cerca de La Palmilla"; Triana 1854:14).A careful examination of the specimen reveals that it consists of three separate fragments, two of which (Fig. 1 A, B) match Triana's (1854) original description of B. ibaguensis, a species characterized, among other traits, by the leaf blades with glabrous lower surface, the hypanthium 3-4(-5) mm long, and the calyx lobes (2-)3-5 mm long, ascending to patent at right angle to hypanthium during anthesis, with entire margins.Only the smaller fragment (Fig. 1B) shows the dilated anther opening that distinguishes the genus Burmeistera.The third fragment (Fig. 1C) clearly differs in the leaf blades with puberulous lower surface, the hypanthium 6-8 mm long, and the calyx lobes > 1 cm long, strongly reflexed, with dentate margins.This third fragment clearly coincides to the protologue and the corresponding illustration of B. rivina (Wimmer 1943, Fig. 33:129), as well as the syntypes (Pennell 10421 and 10323) deposited in GH and NY, both examined by the authors.Additional characters that distinguish B. rivina from B. ibaguensis are the corolla tube 1.5-1.7 cm long and dorsal lobes 0.8-1 cm long; the filament tube puberulous, and the anther tube exserted 1.5-2 cm between the corolla dorsal lobes.The set of characters described above for B. rivina is diagnostic to species level throughout the genus Burmeistera (see e.g., Gleason 1925, Wimmer 1943, Wilbur 1976a, b, Jeppesen 1981, Lammers 1998, Garzón-Venegas & González 2012, Garzón-Venegas et al. 2012, 2013).Thus, floral characters of B. rivina, especially the shape and orientation of calyx lobes and corolla, suggest close affinities to other species with long, strongly reflexed sepals, e.g.B. tenuiflora Donn.Sm., from Central America.
Burmeistera ibaguensis and B. rivina are partially sympatric in the Central Cordillera of Colombia (see below), and it is likely that the specimen was accidentally mixed either in the field or during the shipment and handling of Triana's collections from Bogotá, Colombia to Europe.Díaz-Piedrahíta (1999) described in detail this tortuous route, first to Paris, and then to London.The idiosyncratic method by which Triana numbered his collections could have added further confusion, as they consisted of a first number corresponding to Endlicher's (1841) generic numbers, followed by a personal collection number for each genus.Thus, all Burmeistera specimens collected by Triana bear the number 3069 (after the number given to Centropogon by Endlicher 1841:264), and this particular type specimen can be distinguished by the specimen number 3069.1.1/88.It corresponds to the BM catalogue number 1583.
Because we provide evidence that the type specimen associated with the name Burmeistera (Fig. 1) is mixed, we proceed to designate the fragments labeled as A and B in Fig. 1  x (2.5-)3.5-4.5(-6)cm, distal laminas (2.5-)3-6(-7) x 1-3 cm, membranous, sometimes slightly bullate, base frequently slightly asymmetric, obtuse to round, sometimes truncate, margins slightly sinuose-serrulate especially on the distal third of the lamina, with c. 30 teeth per side in the proximal leaves, and 7-12 teeth per side in the distal leaves, teeth in two size classes, the short ones interspersed with the long ones, obtuse, apex acute to shortly acuminate and often ending in a callose tooth, adaxial surface dark green, glabrous, abaxial surface green suffused with purple, glabrous, venation brochidodromous, prominent on the abaxial surface, (7-)9-11(-14) pairs of secondary veins in the proximal leaves and 5-10 pairs in the distal leaves, irregularly interspaced, higher order venation conspicuous, reticulate, submarginal vein present but interrupted.Flowers solitary, axillary to the distal leaves, often congested at the apex of the branches due to the shortened internodes between young leaves; peduncle 5-6 cm long, 1-2 mm in diameter, ascending, green and often suffused with maroon, glabrous, ebracteate.Hypanthium obconic to subglobose, 3-4(-5) x 3-5(-7) mm, green and often suffused with maroon, glabrous, with 10 conspicuous longitudinal veins; calyx lobes linear to narrowly ovate, (2-)3-5 mm x 0.5-1.5 mm, glabrous, ascending to patent at right angle with respect to the hypanthium during anthesis, secondary veins inconspicuous, margins entire, apex acute, interlobular sinuses 1-4 mm wide; corolla nearly straight during anthesis, 1.5-2.2cm long, glabrous outside, green to green and pale yellow, often suffused with maroon especially around the base, green inside, tube straight, 9-14 x 2-3 mm at its midpoint, base and throat ventricose, lobes erect, with acute apices, slightly unequal in size, dorsal lobes oblong-triangular, falcate, 10-15 x 3-5 mm, lateral lobes oblong-triangular, falcate, 5-7 x 2-3 mm, ventral lobe ovate, deflexed, 4-5 x 4 mm; filament tube 2-2.5 cm long, glabrous, anther tube obconic, 4-5 mm long and 4 mm in distal diameter, exserted 10-15 mm between the corolla dorsal lobes, dorsal and lateral anthers 4-5 mm long, the two ventral anthers 3-4 mm long, glabrous; stigma bilobed, discoidal during post-anthesis and exserted 3-4 mm from the anther tube.Berries subglobose to ovoid, 0.8-1 cm diam., fleshy, not inflated, green to green and pale yellow      of Burmeistera.We also thank the Fundación para la Promoción de la Investigación y la Tecnología, Banco de la República, and the Dirección Nacional de Investigaciones, de la Universidad Nacional de Colombia, for funding the projects 2413/2009 and 201010011272, respectively.Thanks also to J. Betancur (Institute of Natural Sciences, National University of Colombia) and his research team for providing the photographs of Fig. 2, and to N. Pabón-Mora (Universidad de Antioquia) for editing the figures.Finally, we acknowledge three anonymous reviewers, who made important comments and suggestions.

Figure 3 .
Figure 3. Burmeistera ibaguensis (Luteyn 11989, spirit collection at NY). A. Distal portion of a flowering branch.B. Preanthetic flower, ventral view; calyx lobe removed.C. Preanthetic flower, lateral view, a lateral petal removed to show ventral anthers not barbate.D. Longitudinal section of flower at late anthesis, lateral view.E. Detail of ventricose throat and corolla lobes.F. Detail of anther tube that is not barbate and exserted stigma.G. Forming berry, top view.H. Mature berry, lateral view.I, J. Mature berry and subtending leaf in adaxial (I) and abaxial (J) views.Scale bars: 2 mm in A-C, E, F, G; 5 mm in D, H-J.

Figure 4 .
Figure 4. Burmeistera rivina (Murcia UMK-181, HUA); A. Flowering branch.B. Detail of flower and floral buds.C. Berry.Note the long, reflexed sepal lobes in flowers and berries, and their similarity with those of Fig. 1C; note also the similarity of this specimen with Wimmer´s (1943, Fig. 33:129) illustration of this species.Scale bars: 1 cm in A; 5 mm in B, C.
as the lectotype of B. ibaguensis.This is in accordance to the Article 9.14 of the current International Code of Botanical Nomenclature (McNeill et al. 2012), which states that "[W] hen a type (herbarium sheet or equivalent preparation) contains parts belonging to more than one taxon, the name must remain attached to the part that corresponds most nearly with the original description or diagnosis."Additionally, revised descriptions of B. ibaguensis and B. rivina are provided.