Taxonomic history
The genus Orthophytum Beer was first described based on a single specimen collected byJohann Emanuel Pohl in 1820 in a region of the Brazilian state of Minas Gerais which is nowadays known as the Medio Jequitinhonha region. The specimens of the type-species of Orthophytum, (O. glabrum (Mez) Mez) are characterized by being saxicolous and stout plants, without a basal rosette present during anthesis and having a distinct and elongate peduncle.
According to Louzada & Wanderley (2010), when describing the genus, Beer (1854) did not apply any specific epithet to the analysed specimen (Pohl3436). In 1891, Carl Mez, who perhaps did not know about the description of the genus Orthophytum almost 40 years earlier, described Prantleia Mez, as a new genus of Bromeliaceae with two species P. glabra Mez and P. leprosa Mez. However, one of the species placed in Prantleia was described based on Pohl 3435 - the same collection that Beer (1854) used to describe Orthophytum. Consequently, four years later Mez (1896) recognized the first description of Orthophytum and decided to synonymize Prantleia under Orthophytum, presenting two new combinations for the two species.
At the beginning of the 20th century, the botanist Ernst Heinrich Georg Ule described two new genera of Bromeliaceae based on specimens collected during field work in the Brazilian state of Bahia: Cryptanthopsis Ule and Sincoraea Ule (Louzada & Wanderley, 2010).
Cryptanthopsis saxicola Ule was first described based on a tiny bromeliad specimen with a short but evident peduncle, growing on top of granitic rock outcrops in elevations not exceeding 200 m. In contrast, Sincoraea amoena Ule grows on quartzite rock outcrops above 800 m elevation (personal observation) and it is mainly characterized by the complete absence of a peduncle - rendering a sessile inflorescence (Fig. 1).
According to its protologue, Cryptanthopsis is a genus morphologically distinct from Sincoraea, especially by the presence or absence of the peduncle respectively. However, even with the striking differences mentioned above, Smith ( 1940) described C. navioides L.B. Smith as a new species with sessile inflorescences and with the base of the central leaf blades red during the anthesis, similar to Sincoraea amoena.
After that, Smith (1955) recognized both species as belonging to the same genus and proposed the synonymy of Cryptanthopsis navioides and Sincoraea amoena under the genus Orthophytum (Orthophytum navioides (L.B. Sm.) L.B. Sm. and Orthophytum amoenum (Ule) L.B. Sm.). With this decision, Orthophytum now included species characterizing two morphological groups, one with sessile inflorescences and other with pedunculate inflorescences (Smith & Downs, 1979; Wanderley, 1990; Leme, 2004; Wanderley & Conceicao, 2006 ; Louzada & Wanderley, 2010).
After Smith (1955) many other species were described in both morphological groups of Orthophytum and then Leme (2004), in an attempt to organize the species of Orthophytum, called the two morphological groups of the genus the "complex with sessile inforescence" (CSI) and the "complex with scapose (=pedunculate) inflorescence" (CPI). Further, the author, again based on morphological features, recognizes some groups of species within the complexes and calls them subcomplexes. Thus, the CSI includes the subcomplexes "amoenum", "supthutii" and "vagans", and the CPI includes the subcomplexes "disjunctum", "leprosum" and "mello-barretoi".
It is important to highlight that the complexes and subcomplexes of Leme were proposed to provide some guidance for future taxonomic studies, and the inclusion of species in each group was based exclusively on morphological characters without any reference to molecular phylogenetic evidence.
During a study on the species of Orthophytum that have sessile inflorescences, Louzada (2008) argued for the segregation of the species of the subcomplex supthutii into a distinct genus. In this group are included O. supthutii Gross & Barthlott and O. itambensis Versieux & Leme that are morphologically related mainly by the presence of lanceolate petal appendages in contrast to the lacerate/fimbriate petal appendages in the other species of Orthophytum. Based on this morphological character and the evidence that these two species are in a distinct lineage (Schulte et al. 2009) the new genus Lapanthus Louzada & Versieux was proposed to better accommodate these species (Louzada & Versieux, 2010). In the same year a taxonomic revision of Orthophytum with sessile inflorescences was published (Louzada & Wanderley 2010). In this study are included two of the three subcomplexes of CSI of Leme, since the species that belonged to the "supthutii subcomplex" were elevated to the new genus cited above. Still, two groups of species recognized by Leme, the "vagans subcomplex" and "amoenum subcomplex", remained in CSI.
Phylogeny of Orthophytum and related genera
In the most comprehensive phylogeny with focus on Orthophytum species (Louzada et. al.2014) Lemes two complexes appear as non-monophyletic groups. The sub-complex vagans is phylogenetically related to the majority of species with pedunculated inflorescences. On the other hand, the genus Lapanthus is confirmed as a monophyletic group as well as is the subcomplex amoenum. However, the relationship with the other groups was not elucidated due the lack of statistical support in some lineages of the referred phylogeny.
Orthophytum vs. Sincoraea: morphological evidence
After the removal of Lapanthus, there are still two morphological groups based on the presence or absence of a peduncle in the species of Orthophytum. Nevertheless, other features have to be analyzed to improve the understanding of the genus. Molecular evidence suggests the clade with the species of subcomplex vagans is the sister group of CPI. Moreover, some morphological features of species of subcomplex vagans such as petals with obtuse-cucullate apex and cupulate petal appendages are also found in the species of the first lineage diverging from the clade containing all species with pedunculate inflorescences. The species of subcomplex amoenum have an inflorescence morphology distinct from the remaining species. The sessile flowers of this group emerge from the top of a short and inconspicuous stem - resembling an Asteraceae capitulum - while in other species of the genus the flowers are spread along the peduncle or in a long caulescent stem.
Therefore, based on the morphological evidence and the monophyly of the subcomplex amoenum we re-establish the genus Sincoraea for the species with short caulescent habit and sessile inflorescences that are currently included in Orthophytum. In this study, thus, we include an updated description of Sincoraea, besides the new combinations and a new synonymization.
Sincoraea Ule, Bot. Jahrb. Syst. 42:191.1908.
Type:- S. amoena Ule = Orthophytum amoenum (Ule) L.B.Sm. Smithsonian Misc. Collect. 126(1): 33, 179 (1955)
Plants saxicolous, stoloniferous in all populations observed, with inconspicuous short stems covered by the leaf sheaths.
Leaves forming a distinct rosette, straight, arcuate or falcate;
leaf sheaths imbricate, triangular, white-greenish, lepidote to glabrous, margins serrate;
leaf blades coriaceous to subcoriaceous, linear-triangular to narrowly-triangular, flat to concave, lepidote to sparsely lepidote on both surfaces, margins serrate, prickles densely or laxly arranged.
Inflorescence sessile, simple or compound;
primary bracts foliaceous or subfoliaceous.
Floral bracts green or red, margins serrate to serrulate, apex pungent.
Flowers sessile;
sepals free, erect to suberect, asymmetric or rarely symmetric, apex acute, acuminate, mucronate or mucronulate;
petals free, white, spathulate, with two callosities laterally placed to the filaments, margins entire, apex obtuse.
Petal appendages sacciform, lacerate or digitate.
Epigynous tube present or absent.
Stamen included;
flaments filiform, first whorl free, second whorl adnate to the petals;
anthers, dorsifixed obtuse.
Stigma simple-erect.
Fruits ovoid, with persistent sepals.
Seeds ovoid, striate.
Artificial key to the species of Sincoraea
1a Inflorescence simple => 2
1a Inflorescence compound (rarely simple in S. ulei) => 5
2a Sepals without glandular trichomes => S. hatschbachii
2b Sepals with glandular trichomes => 3
3a Sepals narrowly triangular (6:1 or 3:1) => S. humilis
3b Sepals triangular (2:l or 3:2) => 4
4a Leaf blades 2-2.3 x 0.4-0.7 cm; petals c.2.5 cm long; Jacobina-BA => S. navioides
4b Leaf blades 2.5-7 x 0.3-0.4 cm; petals c.2 cm long; Mucugd-BA => S. mucugensis
5a Floral bracts and sepals pink to red => 6
5b Floral bracts and sepals green => 9
6a Leaf blades abaxially densely lepidote (completely covered by scales) => 7
6b Leaf blade abaxially laxly Iepidote => 8
7a Leaf blades adaxially glabrous or sparsely lepidote at the base; petal apex subacute => S. burle-marxii
7b Leaf blades adaxially densely lepidote at the base; petal apex obtuse => S. ulei
8a Leaf blades 0.5-1 cm wide, marginal prickles 1.5-3 mm long; floral bracts c. 1.8 x 1 cm, anthers3.2-3.5 mm long. => S. amoena
8b Leaf blades 0.2-0.4 cm wide; marginal prickles 0.3-0.4 mm long; floral bracts 0.9-1.2 x 0.7; anthers c.2.5 cm long . => S. ophiuroides
9a Leaf blades with a white woolly-lepidote ring surrounding the inflorescence => S. albopicta
9b Leaf blades with a green and glabrous ring surrounding the inflorescence => 10
10a Leaf blades c. 1.5 cm wide, margins densely serrate => S. heleniceae
10b Leaf blades 0.4-07 cm wide, margins laxly serrate => S. rafaelii