Orthocryptanthus (Leme, S. Heller & Zizka) Leme, Zizka & Paule, 2022

Leme, Elton M. C., Zizka, Georg, Souza, Everton Hilo De, Paule, Juraj, De Carvalho, Jordano D. T., Mariath, Jorge E. A., Halbritter, Heidemarie & Ribeiro, Otávio B. C., 2022, New genera and a new species in the “ Cryptanthoid Complex ” (Bromeliaceae: Bromelioideae) based on the morphology of recently discovered species, seed anatomy, and improvements in molecular phylogeny, Phytotaxa 544 (2), pp. 128-170 : 145-148

Orthocryptanthus (Leme, S. Heller & Zizka) Leme, Zizka & Paule , stat. nov.

Basionym:— Orthophytum subg. Orthocryptanthus Leme, S. Heller & Zizka, Phytotaxa 318: 77. 2017.

Type:— Orthophytum vasconcelosianum Leme, Phytotaxa 205: 287. 2015 (holotype RB!, isotype HB!).

Diagnosis:— Orthocryptanthus differs from Orthophytum mainly by petals without appendages (vs. bearing well developed appendages) or sometimes with sacciform appendages (vs. echinatiform), stamens distinctly unequal (vs. equal to subequal), stigma simple-patent or simple-erect (vs. simple-dilated), without papillae or inconspicuously papillate (vs. conspicuously papillate), sepals as long as the fruit to 1.2 times longer (vs.1.3–3 times as long as the fruit), larger seeds (2–3 × 1–2 mm vs. 1.5–2 × 0.6–1 mm), non-angulose (vs. angulose), and seed tegmen with deep undulations or sinuosities (vs. costate).

Description:— Plants saxicolous, monoecious, usually long caulescent or sometimes stemless, propagating by short axillary shoots or stolons. Leaves coriaceous; sheath inconspicuous; blade very narrowly triangular, densely white lepidote at least abaxially to glabrous, usually concolorous at anthesis, sometimes the upper ones around the inflorescence turning reddish but not forming a colored ring, not narrowed at the base, margins straight, spinulose. Inflorescence compound to sometimes pseudosimple, sessile or sometimes inconspicuously pedunculate, corymbose or a compact glomerulate spike, with basal/outer inconspicuously stipitate fascicles; primary bracts foliaceous; basal fascicles 2–9-flowered, flabellate, pulvinate; floral bracts in the fascicles triangular to narrowly triangular, thin in texture, about equaling the midpoint of the sepals. Flowers sessile, 16–24 mm long, odorless or fragrant, diurnal; sepals free, narrowly lanceolate to ovate-lanceolate, acuminate to caudate, not pungent, densely white lepidote to glabrous; petals lanceolate, narrowly obovate or narrowly spathulate, free, 3.5–4.5 times longer than wide, the blades suberect and forming a subtubular corolla to subspreading-recurved and forming an open, fan blade-like corolla at anthesis, partially exposing the stamens, lilac-rose or white, unappendaged or sometimes bearing sacciform appendages, with well developed longitudinal callosities with glandular trichomes. Stamens erect; filament distinctly unequal in length, the antesepalous ones free, the antepetalous ones distinctly shorter than the antesepalous ones, and basally connate with the petals; anther sublinear, distinctly bilobed at the base, not laterally complanate, straight at anthesis, cream; pollen sulcate, oblate, 40–45 μm, reticulate, lumina width of the reticulum decreasing slightly towards the sulcus margins, the sulcus narrow, ornamented with small exine elements predominantly coherent and attached to the sulcus margins, margins weakly defined to indistinct. Ovary usually clavate; epigynous tube inconspicuous; ovules globose, obtuse to obtusely apiculate, few to many; stigma simple-patent or simple-erect, rose-lilac or white, distinctly exceeding the anthers, lobes spreading or suberect, longer than wide, but broadened in the distal part, margins inconspicuously crenulate, without papillae to inconspicuously papillate. Fruits baccate, 4–7 × 3.5–6 mm, sepals persistent, equaling the fruit length to 1.2 times longer; seeds 2–3 × 1–2 mm, 2–8 in number per fruit.

Distribution and habitat:— Orthocryptanthus is endemic to the microregion of Aimorés, in the mesoregion of the Rio Doce valley, Minas Gerais state, occurring in the counties of Alvarenga, Conselheiro Pena, and Santa Rita do Itueto, located in the Atlantic Forest domain, about 400 to 450 km from Belo Horizonte and 45 to 65 km from the border with Espírito Santo state. The species are saxicolous, living on rocky outcrops in open to partially shaded sites, at 1000 to 1441 m elevation.

In the neighboring counties of Alvarenga and Conselheiro Pena, Orthocryptanthus vasconcelosianus (fig. 9 A) and O. arcanus (fig. 9 C) live in a peculiar intermediate vegetation type of Campos Rupestres, which is related to the Cerrado biome, and Campos de Altitude, which is related to the Atlantic Forest biome (see map in Leme et al. 2017b, fig. 4). However, in the county of Santa Rita do Itueto, O. santaritensis (fig. 9 B) grows in a more typical Campo Rupestre vegetation in direct sunlight.

The phylogenetic reconstruction of Schulte & Zizka (2008) and the results obtained by Leme et al. (2017b) and in this study indicated that groups of genera with similar geographic distribution (and possibly origin) might be closer related, often forming monophyletic groups. Similar results were obtained by Aguirre-Santoro et al. (2015, 2016) when analyzing putative disjunct lineages in Ronnbergia E. Morren & André (1874: 120) . The findings of Schulte & Zizka (2008) and the results of Sass & Specht (2010) are confirmed here, since monophyletic clades in the “Cryptanthoid complex” that reveal joint ecological preferences and/or sympatric ranges can in many cases be clearly correlated with morphological characters. This is the case of Orthocryptanthus arcanus , O. santaritensis and O. vasconcelosianus which are microendemic species growing in the same microregion and sharing similar altitude range and ecological conditions. The range of Orthocryptanthus is distinct but comes close to the potential range of Cryptanthus and Orthophytum subg. Orthophytum ( Leme et al. 2017b, figs. 1, 3 ), without documented overlapping, at least not for the isolated mountains they inhabit.

The morphologically unique Krenakanthus roseolilacinus also grows in the same region of occurrence as O. arcanus and O. vasconcelosianus , in the counties of Alvarenga and Conselheiro Pena, but it lives in lower elevated and not sun-exposed sites, in a different ecological niche inside fragments of humid Atlantic Forest, from 255 to 810 m elevation, and presents amazingly distinct morphological features (see below).

Etymology:—According to the protologue ( Leme et al. 2017b), the name Orthocryptanthus is a combination of the Greek “orthós”, which means straight, upright, vertical or erect, like an “erect Cryptanthus ”, in reference to the long caulescent habit of its type species, which was the first one discovered in this group.

Distinctive characters:— Orthocryptanthus is strongly supported as a monophyletic clade (1 PP, 81 BS).In contrast to the previous results obtained by Leme et al. (2017b) of a weakly supported polytomy composed of “ Capixabanthus-Krenakanthus-Orthocryptanthus - Orthophytum duartei clade”, Orthocryptanthus is now strongly supported (1 PP, 100 BS) as a sister of the whole clade Capixabanthus - Clavanthus - Krenakanthus - Orthophytum , which is highly compatible with its geographical and morphological singularity.

It is important to mention that Orthophytum arcanum , now included in Orthocryptanthus , was originally conceived as a member of Orthophytum subg. Capixabanthus , despite it was stated in its protologue that it is “an outlier species without clear morphological affinity with any known species that are endemic to the same region”, mentioning specifically Krenakanthus roseolilacinus , Orthocryptanthus santaritensis , and O. vasconcelosianus ( Leme et al. 2020) . At that time, due to its geographical range, we opted to include it in Capixabanthus, despite of the highlighted presence of discordant features, like the distinct simple-patent stigma type, which, for instance, is in accordance with the stigma type of Orthocryptanthus . In the present study, O. arcanus was recovered with high support (1 PP, 92 BS) in Orthocryptanthus clade, as sister of O. vasconcelosianus which lives in a different mountain (i.e. Pico da Aliança) located in the same microregion of Pico do Padre Angelo, where O. arcanus lives.

Concerning Orthophytum subg. Orthophytum , Orthocryptanthus differs by its petals lacking appendages (vs. basal appendages well developed), or sometimes with sacciform appendages (vs. appendages echinatiform), stamens distinctly unequal in length (vs. equal to subequal), stigma simple-patent or simple-erect (vs. simple-dilated), without papillae or inconspicuously papillate (vs. conspicuously papillate), larger seeds (2–3 × 1–2 mm vs. 1.5–2 × 0.6–1 mm), and seed tegmen with deep undulations or sinuosities (vs. costate). The shared long caulescent habit of O. santaritensis and O. vasconcelosianus with some species of Orthophytum subg. Capixabanthus ( O. pseudovagans , O. vagans , and O. zanonii ) justifies a closer comparison between them as follows: Orthocryptanthus differs from those three species, besides the characteristics mentioned above, by its central leaves usually concolorous or sometimes the inner ones turning reddish but not forming a colorful ring around the inflorescence (vs. basal portion of the central leaves and primary bracts turning reddish or bright red, forming a colorful ring around the inflorescence), petals acute, subspreading-recurved at anthesis and forming an open, fan blade-like corolla at anthesis or petals suberect but not forming a subclavate corolla (vs. obtuse-cucullate, suberect and forming a subclavate, subtubular corolla), and the pollen sulcus with small exine elements which predominantly remain coherent and attached to the sulcus margins (vs. exine elements not attached to the sulcus margins), lumina width of the reticulum decreasing slightly towards the margins (vs. reticulum characteristically formed by broad and flat muri with lumina or faveolae significantly decreasing in size towards the sulcus margins and apices, being almost psilate).

Orthocryptanthus can be easily differentiated from Krenakanthus by distinct morphological features,like coriaceous leaves (vs. leaves in most part chartaceous), leaf blades with margins straight (vs. undulate), flowers 16–25 mm long (vs. 42–55 mm long), petals lanceolate, narrowly obovate to narrowly spathulate (vs. broadly spathulate), without appendages or sometimes with sacciform appendages (vs. bearing cupuliform appendages), blades conspicuously narrower (4–5 mm vs. 15–20 mm wide), remaining erect after anthesis (vs. flaccidescent afterwards), stamens not deeply included and partially visible at anthesis (vs. deeply included and not visible), stigma simple-patent or simpleerect (vs. conduplicate-spiral), persistent sepals equaling the fruit length to 1.2 times longer than the fruit (vs. 2.5–3.5 times longer). Both genera occur in the same region, although Krenakanthus grows in lower elevated sites as mentioned before ( Leme et al. 2017b, fig. 4).

The long caulescent habit of two species of Orthocryptanthus is also observed in some species of Hoplocryptanthus . Besides the distinct geographical range of this new genus (i.e., mesoregion of the Rio Doce valley vs. the Iron Quadrangle and in neighboring areas), it differs from Hoplocryptanthus by having free petals (vs. connate), bearing glandular callosities (vs. glabrous callosities), stamens distinctly unequal in length (vs.equal), filaments not forming a concrescent structure with petals and style (vs. forming such a basal concrescent structure), stigma simple-patent or simple-erect (vs. cylindric-distent), and by larger (2–3 × 1–2 mm vs. 1.5–2 × 1–1.5 mm) and much fewer seeds (2–8 vs. 35–75 in number).

Species:— Orthocryptanthus is composed of three species.

Figures

FIGURE 3. Seed morphology. Seeds in lateral view (A, C–E, G–J, M, O, Q–T) and raphe view (B, F, H, L, N, P). A–B. Cryptanthus aff. beuckeri (Leme 3891). C. C. colnagoi (Leme 9154). D. C. pickelii (Leme 9665). E–F. C. bahianus (Leme 9616). G-I. Siqueiranthus cinereus (Leme 9785). G and I show two morphological variations for this species. J. Krenakanthus roseolilacinus (Leme 8922). K–L. Orthocryptanthus vasconcelosianus (Leme 8673). M–N. Orthophytum maracasense (Leme 9617). O–P. O. compactum (Leme 5604). Q. O. leprosum (Leme 7897). R. O. pseudovagans (Leme 6821). S. O. vagans (Leme 505). T. O. saxicola (Leme 6058). c = chalazal region or inconspicuous chalazal appendage. m = micropylar region. r = raphe. Bars = 0.5 mm.