Nidularium antoineanum, Wawra, 1880

× Nidunelia rodrigoana Leme, nothogen. et nothosp. nov. ( Figs. 23 A–D View FIGURE 23 , 24 A–J, W–Y View FIGURE 24 )

Diagnosis:—This new nothogenus and nothospecies is morphologically intermediate to its putative parents, Nidularium antoineanum and Quesnelia lateralis Wawra (1880b: 149) , differing from them by the combination of compound inflorescence with involucral primary bracts, fascicles elongate, petals shortly connate at the base, bearing well-developed longitudinal callosities as well as irregularly long laciniate appendages at the base, and stigma irregularly spiraled, bluish-white.

Type:— Brazil. Rio de Janeiro: Teresópolis , P . E . Três Picos , 1,650 m elev., October 2021, R . F . da Silva s.n., cult. E . Leme 10111 (holotype RB!) .

Discription:— Plants terrestrial, ca. 50 cm tall when in bloom, propagating by short basal shoots. Leaves 10–14 in number, subcoriaceous, forming a funnelform rosette; sheath elliptic to narrowly elliptic, 13–16 × 7–8, whitish near the base and greenish toward the distal end, subdensely and inconspicuously brown lepidote toward the base adaxially, sparsely and minutely white lepidote toward the distal end, finally nerved; blade lingulate, narrowed toward the base, 37–40 × 4.3–5.3 cm, subdensely and inconspicuously white lepidote abaxially, sparsely and inconspicuously white lepidote adaxially, green, finely nerved, apex acute and apiculate, margins densely spinose, spines triangular, antrorse, 0.3–0.5 mm long, 2–6 mm apart; peduncle erect to suberect, 30 × 0.7–0.8 cm, green, glabrous, smooth; peduncle bracts the basal ones foliaceous, the distal ones resembling the basal primary bracts, green, equally arranged along the peduncle, erect, distinctly exceeding the internodes and completely covering the peduncle. Inflorescence (fertile part) shortly corymbose, narrowly obconic, twice branched, distinctly elevated above the rosette but shorter than the leaf blades, 6.5–7.5 cm long (including the primary bracts), ca. 4.5 cm in diameter at the apex; primary bracts resembling the involucral bracts, but gradually smaller, the outer ones narrowly ovate-lanceolate, apex acute and apiculate, suberect to nearly erect, ca. 9 × 3.5 cm, subdensely but inconspicuously white lepidote, green toward the base and reddish toward the margins and apex to completely dark red, densely spinose toward the apex, spines antrorse, ca. 0.5 mm long, 3–5 mm apart; primary fascicles ca. 45 × 15–17 mm, complanate, slightly elongated, ca. 4-flowered, distinctly stipitate, stipe trapeziform, ca. 5 × 7 mm, greenish-white, glabrous, slightly corrugate; floral bracts slightly exceeding the sepals, carinate, subtriangular to ovate, pale rose toward the apex, 19–20 × 9–13 mm, finely nervate, membranaceous, inconspicuously lepidote toward the apex to glabrous, apex acute and apiculate, margins entire. Flowers ca. 30 mm long, sessile, diurnal, odorless; sepals narrowly elliptic-lanceolate to suboblong, symmetrical, apex acute, 11–12 × 3.5 mm, connate at the base for 3–3.5 mm, glabrescent, greenish, finely nerved, ecarinate, thin in texture; petals lingulate, 13–20 × 3.5 mm, shortly connate at the base, membranaceous, apex obtuse-cucullate, greenish-white near the base, cobalt-blue toward the apex, erect and forming a tubular, subclavate corolla, bearing at the base 2 conspicuous callosities about equaling the filaments, as well as 2 basal appendages; appendages 1.5–2 × 1 mm, irregular in shape, more or less cuneate, irregularly long laciniate; filaments 8–13 mm long, the antepetalous ones adnate to petals for the basal half of their length, the antesepalous ones shortly adnate to the petal tube; anthers narrowly suboblong, ca. 5 mm long, base obtuse, apex apiculate, fixed near the middle; pollen narrowly ellipsoid, ca. 40 μm in diameter, porate, pores medium, exine broadly reticulate, lumina irregularly polygonal; style blue; stigma conduplicate-spiral, irregularly spiraled, ca. 2 × 1 mm, lobes with minutely crenulate margins, without papillae, bluish-white; ovary oblong to subquadrate, trigonous, 10–12 × 5–5.5 mm, white, glabrous; epigynous tube ca. 1.5 mm long; placentation from median to apical; ovules many, subcylindrical, obtuse. Fruits unknown.

Distribution and habitat:––× Nidunelia rodrigoana was found as terrestrial in a nebular Atlantic Forest, about 1,650 m elevation, inside the Três Picos State Park, Teresópolis county, in the Serra dos Orgãos mountain range, Rio de Janeiro state, southeastern Brazil (fig. 23A). It forms small and dense groups of plants (fig. 23 C), growing sympatrically with the ornithophilous Nidularium antoineanum Wawra (1880a: 113) (fig. 23 E) and Quesnelia lateralis (fig, 23 F), which were found as terrestrials, saxicoles and epiphytes surrounding the small clumps of this new nothespecies.

Etymology:––The name of this new nothogenus is a combination of the names of its putative parents, Nidularium Lemaire (1854: 60) and Quesnelia Gaudichaud (1842: t. 54). The new nothospecies, × Nidunelia rodrigoana, honors its collector, the researcher of the Três Picos State Park, Rodrigo Freitas da Silva, currently studying the flora of the most elevated mountain parts of this amazing protected area. The attention of Mr. Silva was attracted to this new nothospecies by its unusual general appearance completely distinct from anything he had ever seen or documented so far for this conservation unit.

Distinctive characters:—This new nothogenus and nothospecies is morphologically intermediate between its putative relatives, Nidularium antoineanum Wawra (1880a: 113) and Quesnelia lateralis Wawra (1880b: 149) (see table 1), differing from them by the combination of compound inflorescence with involucral primary bracts, fascicles elongate, petals shortly connate at the base, forming a subclavate corolla, bearing well-developed longitudinal callosities, as well as basal irregularly long laciniate appendages, and by the stigma irregularly spiraled, bluish-white.

The pollen of this new species shares the same general characteristics of the pollen of its putative parents, being narrowly ellipsoidal, biporate with large pores, with broadly reticulate exine. In relation to N. antoineanum , this new nothogenus resembles it by the subcoriaceous leaf blades which are narrowed toward the base, the compound, corymbose inflorescence with involucral bracts, the flattened fascicles, petals with obtuse-cucullate apex, and the bluish-white stigma. Its elongate flower fascicles (due to the more elongated inflorescence of Q. lateralis ), the shortly connate petals with well-developed longitudinal callosities, the presence of long-laciniate basal appendages, the filaments shortly adnate to the petals, and the bluish-white color of the stigma with weakly spiraled lobes make it morphologically related to Q. lateralis (table 1).

Natural hybridization is a widespread, crucial phenomenon in plant evolution and diversity ( Soltis & Soltis 2009, Marques et al. 2019) and has been commonly reported mainly involving sympatric, closely related species and can reflect recent divergence time ( Zanella et al. 2016, Neri et al. 2017). In Bromeliaceae , interspecific natural hybridization is quite frequent, as well as in cultivation, and the same can also be said in relation to crosses between genera under controlled conditions, with several ex-situ intergeneric crosses registered ( Smith 1983, Grant & Zijlstra 1998, Lawn, cont. updated). However, bigeneric hybridization is seldom reported for the wild, with only few crosses documented, like × Guzlandia barbieae ( Rauh, 1985: 46) Gouda (2019: 1988), × Hohenmea itapuana B.R. Silva & L.F. Sousa ( Sousa et al. 2003: 73) (see Gouda et al. cont. updated), and three × Niduregelia nothospecies previously published as species ( Leme 2000). Once having overcome the prezygotic barriers, the rarity of the phenomenon in situ may be the consequence of different events during pollen tube growth, such as the irregular deposition of callose in pollen tubes, entangled pollen tubes in the style, and arrest of pollen tube growth in the style, among other causes ( Souza 2013).

Despite Nidularium antoineanum and Quesnelia lateralis share sympatric occurrence and have similar flower dimensions, pollen morphology, and ornithophilous pollination syndrome, they are morphologically unrelated species and also likely distant phylogenetic relatives ( Bratzel et al. 2023), which highlights the importance of this new, well field-documented nothogenus and nothospecies.

s.n.) and Quesnelia lateralis (Leme 2366) .