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The continuous refinement of the morphological, anatomical, and molecular studies on the genera of the “Cryptanthoid complex”, mainly considering the recently discovered outlier species, gave rise to new interpretations of species relationships. It reinforced the monophyly of the groups, and allowed the recognition of the new monotypic genus Siqueiranthus based on a morphologically unique endemic species from northeastern Brazil on the brink of extinction, as well as generic status for Krenakanthus and Orthocryptanthus previously conceived as subgenera of Orthophytum, with readjustments in the positioning of some species. An integrative approach with the combination of multiple traits and the critical assessment of the diagnostic characters proved to be useful for the establishment of the new monophyletic genera. A new unusual species of Orthophytum is also described and illustrated, and its phylogenetic relationships are discussed.

Krenakanthus (Leme, S. Heller & Zizka) Leme, Zizka & Paule, stat. nov . Basionym:— Orthophytum subg. Krenakanthus Leme, S. Heller & Zizka, Phytotaxa 318: 75. 2017. Type:— Orthophytum roseolilacinum Leme, Phytotaxa 205: 284. 2015 (holotype RB!). Diagnosis:—This new genus differs from Orthophytum by leaves in most part chartaceous (vs. coriaceous or subcoriaceous), leaf blades glabrescent or glabrous (vs. usually distinctly lepidote at least abaxially) with undulate margins (vs. not undulate), flowers 4.2–5.5 cm long (vs. usually to 3 cm long), fragrant (vs. usually odorless), sepals connate (vs. free), petals broadly spathulate (vs. linearlanceolate to narrowly spathulate), spreading at anthesis (vs. erect to suberect with recurved apex only) and flaccidescent afterwards (vs. remaining erect), rose-lilac to lilac-purple toward the apex (vs. usually white or green), petal blades suborbicular (vs. ovate, obovate or elliptic), petal appendages thick, cupuliform (vs. echinatiform or sacciform, rarely cupuliform), stamens deeply included and not visible (vs. not deeply included, partially to distinctly visible), conduplicate-spiral stigma (vs. simple-dilated), and seeds laterally flattened (vs. not laterally flattened), with narrowly oblong raphe in cross section (vs. triangular in cross section). Distribution and habitat:— Krenakanthus roseolilacinus, the single species in the genus, is only known from the type region, in the counties of Alvarenga and Conselheiro Pena, in the microregion of Aimorés which belongs to the mesoregion of the Rio Doce valley, Minas Gerais state (see Leme et al. 2017b, fig. 4). Plants are terrestrial, forming large and dense groups (fig. 8 A) in organic-rich, shallow soils accumulated on rocky surfaces in shady spots inside fragments of humid Atlantic Forest, from 255 to 810 m elevation (see Leme et al. 2017b, fig. 15 A). Krenakanthus grows in the same region of two species of Orthocryptanthus (O. arcanus and O. vasconcelosianus), in the counties of Alvarenga and Conselheiro Pena, but it lives at lower elevated sites (i.e., 255 to 810 m elevation) as a typical Atlantic Forest dweller, not in sun-exposed sites associated to altitude fields/rupestral fields above 1000 m elevation, like O. arcanus and O. vasconcelosianus. It presents amazingly distinct morphological features from them (see respective item below). As microendemic, with a reduced number of individuals in its known subpopulations and living in depleted and strongly fragmented Atlantic Forest habitat, it is considered a critically endangered (CR) species on the basis of the criteria “B1a” and “B2a” of IUCN (2020). Etymology:—According to its protologue (Leme et al. 2017b), Krenakanthus was named to honor the last descendants of the indigenous people native to eastern Brazil (pejoratively designated as “Botocudo” by the European settlers), known for their bravery, their comparatively taller and stronger physique, and by the wooden disks worn to expand their lips and ears. They were typical inhabitants of the Atlantic Forest of the states of Bahia and Espírito Santo, and the “Tupi” people called them “Aimoré” (Pessoa 2009). After being decimated over the centuries, only the “Krenak” group survived, retaining their original language (“Borum”) and living between the cities of Conselheiro Pena and Resplendor, in Minas Gerais (Seki 1992). This new genus was found in the county of Conselheiro Pena, not far from the territory of the remaining “Krenak”. Its single known species is comparatively tall, like the “Krenak”, with unique broad petals resembling expanded lips: with the Greek suffix “anthos”, Krenakanthus means the flower of the “Krenak” people. Distinctive characters:—Positioned in the strongly supported (1 PP, 100 BS) clade “ Orthophytum s.l. ”, K. roseolilacinus is moderately supported (0.94PP) as sister of the remaining“ Orthophytum s.l. ” clade(i.e., Capixabanthus-Clavanthus-Orthophytum). It is a unique species without closer morphological affinity to any known species. Its general vegetative appearance characterized by the combination of long caulescent habit, and comparatively thin-textured leaves with undulate margins give it a very distinct appearance similar as some species of Rokautskyia which are not at all morphologically related to it. It also shares the caulescent habit and sessile inflorescence with three species of Orthophytum subg. Capixabanthus [i.e., O. pseudovagans, O. vagans, and O. zanonii Leme (2004: 72)]. However, the caulescent habit is not an exclusive character of any genus, despite being important to characterize a group of species and segregating them. In fact, the flower morphology of Krenakanthus is conspicuously distinct from Orthophytum subg. Capixabanthus in all aspects, except for the similar cupuliform petal appendages. Despite other differences in flower features, its thick, cupuliform petal appendages are also observed in O. foliosum Smith (1941: 58), O. rubiginosum Leme (2005: 158), and O. zanonii, while the sacciform type is noticed in O. compactum Smith (1966: 462) and O. vagans. Orthophytum pseudovagans has both cupuliform and sacciform petal appendage types. It is important to highlight the unique combination of flower, fruit, and seed morphology and anatomy features of K. roseolilacinus, which distinguishes it from all other “Cryptanthoid” genera, such as sepals distinctly connate, petals with a very narrow proximal portion in contrast with the much broader spreading blade (fig. 8 D), the deeply included stamens not visible at anthesis (figs. 8 B, E), the conduplicate-spiral stigma type (fig. 15 J), fruits obtusely angulose, yellowish toward the apex, and the long and narrow (2–3 × 0.7–1 mm), slightly to distinctly curved seeds (fig. 18 O), which are laterally flattened, bearing fleshy, narrowly oblong raphe in cross-section (see item discussion above), not detected in any other Cryptanthoid species. In general, Krenakanthus differs from Orthophytum subgg. Orthophytum, Capixabanthus, and Clavanthus by having leaves that are for the most part chartaceous (vs. coriaceous or suboriaceous), leaf blades glabrescent or glabrous (vs. usually distinctly lepidote at least abaxially) with undulate margins (vs. not undulate), inflorescence sessile (vs. usually on a distinct peduncle, except for three species of Orthophytum subg. Capixabanthus mentioned above, and members of Sincoraea), flowers 42–55 mm long [vs. usually to 30 mm long, except for the outlier O. macroflorum Leme & M. Machado (2005: 171) with 50–65 mm long flowers], fragrant (vs. scentless), sepals connate (vs. free), petals broadly spathulate (vs. linear-lanceolate to narrowly spathulate), spreading at anthesis (vs. erect to suberect with recurved apex) and flaccidescent afterwards (vs. remaining erect afterwards), rose-lilac to lilac-purple toward the apex (vs. usually white or green), petal blades suborbicular (vs. ovate, obovate or elliptic), petal appendages cupuliform (vs. echinatiform, sacciform or rarely cupuliform in Orthophytum subg. Capixabanthus), stamens deeply included and not visible (vs. not deeply included, partially to distinctly visible), and the conduplicate-spiral stigma (vs. simple-dilated, simple-erect or simple-patent). Despite not morphologically related to Krenakanthus, Rokautskyia reveals a highly supported sister position in relation to the whole “ Orthophytum s.l. ” clade (i.e. Capixabanthus-Clavanthus-Krenakanthus-Orthocryptanthus-Orthophytum) in which the two accessions of Krenakanthus from distinct subpopulations are included. Similarities are fragrant flowers and the broadly spathulate petals with suborbicular blades spreading at anthesis, and the shaded Atlantic Forest habitats. Differences are striking such as the rose-lilac to lilac-purple petals (vs. white) with cupuliform appendages (vs. exappendiculate), the stamens deeply included and not visible (vs. not deeply included and at least partially visible), and the conduplicate-spiral stigma blade (vs. simple-imbricate). A full description of Krenakanthus roseolinacinus, here provided as a descriptio generico-specifica [Shenzhen Code, art. 38.5; Turland et al. (2018)], is presented in the sequence and the differences from Orthocryptanthus are discussed in the respective item below Species: — Krenakanthus is a monotypic genus., Published as part of 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, pp. 128-170 in Phytotaxa 544 (2) on pages 142-143, DOI: 10.11646/phytotaxa.544.2.2, http://zenodo.org/record/6503832, {"references":["Leme, E. M. C. Heller, S., Zizka, G. & Halbritter, H. (2017 b) New circumscription of Cryptanthus and new Cryptanthoid genera and subgenera (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular phylogeny. Phytotaxa 318: 1 - 88. https: // doi. org / 10.11646 / phytotaxa. 318.1.1","IUCN (2020) The IUCN Red List of Threatened Species. Version 2020 [online]. Available from: http: // www. iucnredlist. org (Accessed 4 October 2021).","Pessoa, K. N. (2009) Para um estudo da fonologia Krenak: alguns problemas e analise preliminar. Estudos Linguisticos 38: 233 - 247.","Seki, L. (1992) Notas para a historia dos Botocudos (Borum). Boletim do Museu do Indio 3: 1 - 20.","Leme, E. M. C. (2004) Studies on Orthophytum - Part II: two new scapeless species. Journal of the Bromeliad Society 54: 66 - 74.","Smith, L. B. (1941) Bromeliaceas novas ou interessantes do Brasil - I. Arquivos de Botanica do Estado de Sao Paulo 1: 53 - 73.","Smith, L. B. (1966) Notes on Bromeliaceae, XXIV. Phytologia 13: 454 - 465. https: // doi. org / 10.5962 / bhl. part. 21852","Turland, N. J., Wiersema, J. H., Barrie, F. R., Greuter, W., Hawksworth, D. L., Herendeen, P. S., Knapp, S., Kusber, W. - H., Li, D. - Z., Marhold, K., May, T. W., McNeill, J., Monro, A. M., Prado, J., Price, M. J. & Smith, G. F. (eds.) (2018) International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashutten: Koeltz Botanical Books. https: // doi. org / 10.12705 / Code. 2018"]}

Siqueiranthus Leme, Zizka, E.H. Souza & Paule, gen. nov. Diagnosis:—This new genus differs from Cryptanthus by its sepals shortly connate at the base for 1/5–1/3 of their length (vs. connate for 1/2–2/3 of their length), petals free or subfree (vs. basally connate for 1/7–1/3 of their length), appendaged (vs. unappendaged but sometimes bearing well-developed lateral callosities), erect except for the suberect apex and forming a prevailingly tubular corolla (vs. arcuate-recurved at anthesis and forming an open, fan blade-like corolla), stamens unequal in length (vs. equal), anthers basifixed or subbasifixed (vs. dorsifixed), pollen with reticulum showing slightly flat muri and broader lumina (vs. muri typically thick with narrower lumina), sulcus covered with exine elements partially forming a net (vs. sulcus completely covered with exine elements forming a net), stigma simple-erect with tendency to simple-patent (vs. conduplicate patent), epigynous tube well-developed (vs. usually inconspicuous or lacking, rarely distinct), fruits longer and narrower [18–25 × 7–8 (–9.2) mm vs. 12–20 × 9–12 mm], seeds smaller [2–3.5 (–4) × 1.5–3 mm vs. 3.5–5 × 2.5–4 mm], with small and underdeveloped embryo (vs. differentiated fully developed embryo). Type:— Cryptanthus cinereus D.M.C. Ferreira & Louzada, Systematic Botany 45: 460. 2020 [≡ Siqueiranthus cinereus (D.M.C. Ferreira & Louzada) Leme, Zizka, E.H. Souza & Paule]. Distribution and habitat:—In the “Cryptanthoid complex” the geographical distribution proved to be important to strengthen taxa circumscription and clarify relationships, since monophyletic clades that reveal joint ecological preferences and/or sympatric ranges can in many cases be clearly correlated with morphological characters (Leme et al. 2017b). Some of the obtained monophyletic clades often consist of species with similar ranges (and possibly origin), at least at the scale of co-occurrence within a given geographical area, e.g., a specific mountain chain. The known population of Siqueiranthus in the county of Ibateguara, Alagoas state, northeastern Brazil, is situated at the confluence of the ranges of Cryptanthus, which prevails along coastal areas in Atlantic Forest related habitats, and Orthophytum with a more inland geographical distribution, more often associated with open, rocky outcrop environments, in Northeastern Brazil, north of São Francisco river (fig. 10). However, no sympatric occurrence was observed within any species of these three genera, despite the current strong habitat fragmentation and biological impoverishment observed at the type locality of Siqueiranthus just reveal a pale scenario of the local diversity in the present days. The reported confluent occurrence in theAtlantic Forest domain is clearly in accordance with the close morphological relationship of Siqueiranthus and Cryptanthus ‒ it is part of the earliest branching “Cryptanthoid clade” and sister to Cryptanthus itself (fig. 1), and also in some aspect it shares with Orthophytum a given number of vegetative and floral characters (see topic “distinctive characters” below). On one hand, there is no correspondence between the range of Siqueiranthus and the geographic pattern of any other Cryptanthoid genus or subgenus (see distribution maps in Leme et al. 2017b, figs. 1–4). On the other hand, it was already noted that the distribution of Cryptanthus species seldomly overlaps with that of the remaining closely related Cryptanthoid genera (Leme et al. 2017b). Siqueiranthus is microendemic, known only from a small population at the type locality, in the county of Ibateguara. This place belongs to the microregion of “Serrana dos Quilombos”, east of Alagoas state, known to have one of the lowest temperatures in northeastern Brazil. The area belongs to a biogeographic unit, along with areas in the states of Pernambuco, Paraíba, and even Rio Grande do Norte, Ceará, and Piauí, known to be an important South American center of endemism, a “hotspot’s hotspot”, the so called “ Pernambuco Endemism Center”, or simply “Centro Pernambuco ” (Tabarelli & Santos 2004, Silva & Casteleti 2005). The county of Ibateguara is one of the sites of the “Centro Pernambuco ” that have been tagged as epicenter of imminent massive species extinction (Ricketts et al. 2005, Leme & Siqueira-Filho 2007). For instance, the endemic Aechmea marginalis Leme & J.A. Siqueira (2001: 149), also found originally in an Atlantic Forest fragment in Ibateguara two decades ago, was never recollected despite of several attempts to locate it (Leme & Siqueira-Filho 2006), which gives a real perspective of what will happen to Siqueiranthus in a very near future in a clear ongoing scenario of mass extinction, as reported by Pontes et al. (2016). Etymology:—This new genus honors the biologist Prof. Dr. José Alves Siqueira Filho, of the Universidade Federal do Vale do São Francisco – UNIVASF, in Pernambuco, Northeastern Brazil, founder and Curator of the Herbarium HVASF, founder and executive director of the Centro de Referência para a Recuperação em Áreas Degradadas – CRAD, for Caatinga biome. Besides his expertise in taxonomy of Bromeliaceae, reproductive biology, and conservation of the Atlantic Forest and Caatinga biomes, and its flora and fauna, Dr. Siqueira has greatly contributed to the knowledge on the biodiversity of the states of Pernambuco, Alagoas, and neighboring regions in northeastern Brazil, and on the countless critically endangered species by means of his books, scientific articles, and lectures. Distinctive characters:— Siqueiranthus has a general aspect that recalls a member of Orthophytum due to its habit, leaf conformation, texture, trichome coverage, and corolla structure. However, its andromonoecy, pollen, fruits and seeds clearly demonstrate its close relationship with Cryptanthus, justifying the original position of the taxon within this genus proposed by Ferreira & Louzada (2020). This close relationship is also highly supported by molecular data obtained here and reflected by its sister position to the whole clade of Cryptanthus. On the other hand, misinterpretation/overlooking of some important morphological characteristics indicates the need for a reassessment of the combination of its diagnostic features and species delimitation in comparison to Cryptanthus. Ferreira & Louzada (2020) stated in the prolologue that C. cinereus propagates by axillary shoots and rhizomes 5–99 cm long, and that the propagation type is not a consistent character because it may vary within the same species, citing the synonym C. burle-marxii Leme ‒ originally conceived as a long-stolonifeous species ‒, in relation to C. zonatus (Visiani, 1847: 4) Visiani (1854: 9) ‒ typically propagating by axillary shoots ‒ as previously proposed by Ferreira (2016) and also in Ferreira et al. (2021). These authors might have misinterpreted the long and thick stem of C. cinereus as rhizomes because the stem in this species may lay on the floor with age, mainly the proximal older portion, and generates some axillary shoots. Undoubtely, C. cinereus does not produce any rhizome or stolons, which raises questions about the denied importance of propagation type in species delimitation in this group. Concerning flower morphology, it was not noted that prevailingly tubular corolla of C. cinereus is not observed in any Cryptanthus species, but characterizes other Cryptanthoid genera, like Lapanthus and Orthophytum. For instance, the thick petal appendages of C. cinereus, which are scutelliform with tendency to cupuliform, was interpreted by Ferreira & Louzada (2020) as conspicuous callosities, as well as its simple-erect (with tendency to simple-patent) stigma type ‒ a kind of stigma not present in Cryptanthus ‒ was considered to be conduplicate-patent (typical of Cryptanthus and Lapanthus), minimizing the importance of useful morphological differences of this unusual taxon from all other Cryptanthoid members. The highly supported sister position of Siqueiranthus to Cryptanthus in the phylogeny together with the number of distinctive morphological and anatomical characters form the basis for our decision to recognize it as a distinct new monotypic genus. A full description of the species is presented below based on living specimens that flowered at the type locality. In the “Cryptanthoid complex” the presence of andromonoecy in Siqueiranthus is only shared with Cryptanthus, which immediately points towards their close parentage. However, the new genus differs from Cryptanthus by its distinctly caulescent habit (vs. usually stemless or when caulescent stem to 60 cm tall). The caulescent habit, however, is also observed in Hoplocryptanthus, Krenakanthus, Orthocryptanthus, Orthophytum subg. Capixabanthus, and Rokautskyia, despite Siqueiranthus has the longest (to 160 cm long) and thickest (to 2.8 cm in diameter) stem ever documented in the group (figs. 12 E, F). Besides, its sepals are shortly connate at the base for 1/5–1/3 of their length (fig. 13 E) when compared to the sepals of Cryptanthus, which are connate for 1/2–2/3 of their length. On the other hand, petals are free or subfree (fig. 13 F) as in Krenakanthus, Lapanthus, Orthocryptanthus, Orthophytum subgg. Orthophytum, Capixabanthus, and Clavanthus, and Sincoraea. In contrast, petals in Cryptanthus are basally connate for 1/7–1/3 of their length (fig. 13 I). Petals of Siqueiranthus are erect except for the suberect apex at anthesis (figs. 13 C, D) and form a prevailingly tubular corolla [type II, according to Leme et al. (2017b)], like the corolla architecture of Lapanthus and Orthophytum (figs. 13 J, K), while Cryptanthus has an open, fan blade-like corolla of type I (cf. Leme et al. 2017b) formed by arcuate-recurved petals (figs. 13 G, H). It is important to highlight that petal appendages are scutelliform in this new genus (figs. 14 A–C), as in Orthophytum subg. Clavanthus, with a tendency to cupuliform as also observed in Krenakanthus (fig. 14 H) and Orthophytum subg. Capixabanthus. In contrast, Cryptanthus has unappendaged petals, sometimes with well developed lateral callosities (figs. 14 D, E). Filaments in this new genus are unequal in length (vs. equal in Cryptanthus), similar to Clavanthus, Krenakanthus, Orthocryptanthus, and Orthophytum. Anthers have a narrowly subtriangular shape before and at anthesis, with a pronounced bilobed base and an obtuse and inconspicuously apiculate apex (figs. 16 A–D). Size varies greatly before anthesis (to 5 mm long) and afterwards (2.5–3 mm long) due to dehydration during dehiscence, as usually observed in all members of the “Cryptanthoid complex” (see fig. 16). Anthers in Siqueiranthus are considered basifixed or at least subbasifixed because filaments are fixed right at the base of the connective which distinctly separates the two thecae for 3/4 of their length (figs. 16 A, C). Despite being similar, the anthers of Cryptanthus are dorsifixed (figs. 16 F, I) or at least dorsifixed near the base but not basifixed (figs. 16 K, L) like in other members of the “Cryptanthoid complex”. Dehiscence is adaxial similarly to Cryptanthus (anthers not laterally compressed at anthesis), contrasting with the prevailing lateral dehiscence of the anthers of Orthophytum that become laterally compressed at anthesis (figs. 16 N, P). In general, the morphology of the anthers has been neglected in the taxonomy of Bromeliaceae due to the lack of a detailed characterization in species/genera descriptions. However, in a pioneering study, Gardner (1986) associated filament and anther characteristics as the basis for the recognition of five morphological groups in Tillandsia. Later on, Leme et al. (2017a) used anther dehiscence (introrse or latrorse) and the connective exposure during dehiscence to distinguish species of the genera Waltillia Leme et al. (2017a: 29) and Vriesea Lindley (1843: 11), with the anther dehiscence introrse, from Alcantarea (É. Morren ex Mez) Harms (1929b: 802), with anther dehiscence latrorse. In a morphoanatomical study of some species of Dyckia Schultes & Schultes f. (1830: 1194), Carvalho et al. (2016) and Carvalho et al. (2017) highlighted the importance of anther characteristics in species delimitation, analysing, e.g. connective structure and thickening, anther dehiscence, and stamen morphometry. Recently, Büneker (2021) provided detailed illustration and descriptive morphological data of anthers of a complex of Dyckia species, reinforcing the utility of shape, dimension, apex structure, level of development, and influence of the connective in anther dehiscence in circumscribing species and group/complex of species. In Bromelioideae, Leme et al. (2021) adopted length-width ratio of the anthers and the point of filament fixation to distinguish species in Hylaeaicum (Ule ex Mez, 1934: 40) Leme et al. (2021: 45), as well as documented an unusual prolongation structure at the distal end in one species. However, the use of anther characteristics in taxonomy in species/genera delimitatons greatly depends on the accumulation of detailed data, which is still rudimental, and remains challenging. Pollen in Siqueiranthus is smaller when comparared to Cryptanthus (ca. 55 µm vs. 60–70 μm), bearing a reticulum with slightly flat muri and broad lumina (vs. muri typically thick with narrower lumina), sulcus covered with exine elements partially forming a net (vs. sulcus completely covered with exine elements forming a net), somewhat resembling pollen of Krenakanthus and Orthophytum subg. Capixabanthus (see fig. 12). The key-importance of stigma morphology in generic delimitations and the recognition of monophyletic lineages was widely explored by Barfuss et al. (2016) and Leme et al. (2017b), but the existence of several intermediate structures and subtypes makes the use of stigma morphology in Bromeliaceae taxonomy challenging, requiring constant reevaluation mainly when the optimal condition for study is achieved. The precise interpretation of stigma morphology requires caution and some experience because the structures change during development (Varadarajan & Brown 1988): compaction, lobe position, interwining, twisting and spiral folding and length, as well as lobe and margin curvature and presence of papillae (Brown & Gilmartin 1989). Thus, determination of stigma type should ideally be based on flowers immediately before anthesis (Brown & Gilmartin 1989), as was done here to identify the stigma type of Siqueiranthus as simple-erect with a tendency to simple-patent (figs. 15 D–F), differing from the conduplicate-patent stigma of Cryptanthus (figs. 15 A–C), which is also observed in Lapanthus. Simple-erect and simple-patent types were reported by Leme et al. (2017b) for Forzzaea, Orthocryptanthus (figs. 15 G–I), and Sincoraea. The epigynous tube of Siqueiranthus is amazingly well-developed (fig. 18 A), varying from 5 to 8 mm long, while members of the “Cryptanthoid complex” usually lack an epigynous tube or present an inconspicuous one, rarely a distinct epigynous tube is found. However, the reexamination of some Cryptanthus species revealed a larger epigynous tube than previously reported by Leme et al. (2017b), but no longer than 3 mm (figs. 18 B–D). Fruit size, shape, color, fragrance, and calyx length, and persistency proved to be important in species and genera circumscriptions and delimitations in Bromelioideae (Leme 1997, 1998, 2000, Leme et al. 2017b, 2021), and the amazing fruit diversity in this subfamily was for the first time documented in detail by Leme et al. (2021). The large fruits of Siqueiranthus reinforce its close relationship with Cryptanthus, but a closer analysis reveals it has longer and narrower fruits (figs. 18 E, F) than Cryptanthus [18–25 × 7–8 (–9.2) mm vs. 12–20 × 9–12 mm (figs. 18 G–J)], despite some overlapping measures (value in parenthesis indicated in the protologue but not observed in the specimens we studied). In fruit, the sepals are persistent or the distal portion decays in Siqueiranthus (figs 18 E, F), similarly to Cryptanthus (figs 18 G–J), with remnants 6–7 times shorter than the fruit length. Differences in relation to Cryptanthus in this aspect are subtle, because their sepals always decay and the remnants are not more than 2–4 times shorter than the fruit length. The first attempt to systematically use seed morphology on a large scale in Bromelioideae to reinforce genera cincumscriptions was done by Leme et al. (2017b, 2021). They adopted size, shape, number of seeds per fruit, coverage of an aril-like, whitish-translucent jelly substance, and presence of chalazal/micropylar monocaudate or bicaudate appendages as distinctive characters. In Siqueiranthus, seeds are few (10–20 per fruit) and considered large [2-3.5 (-4) x 1.5-3 mm] compared to the seeds of other genera of the “Cryptanthoid complex”, except Cryptanthus for which even larger seeds are reported (3.5–5 × 2.5–4 mm) (see fig. 13). The morphology and anatomy, however, revealed striking differences of the seeds of Siqueiranthus in relation to Cryptanthus from one side, as well as demonstrated important similarities with the seeds of Orthophytum from the other, like its underdeveloped embryo. The observed differences are fully addressed in the item discussion above. A full redescription of Siqueiranthus cinereus, here provided as a descriptio generico-specifica [Shenzhen Code, art. 38.5; Turland et al. (2018)], is presented below. Species:— Siqueiranthus is monotypic., Published as part of 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, pp. 128-170 in Phytotaxa 544 (2) on pages 149-159, DOI: 10.11646/phytotaxa.544.2.2, http://zenodo.org/record/6503832, {"references":["Leme, E. M. C. Heller, S., Zizka, G. & Halbritter, H. (2017 b) New circumscription of Cryptanthus and new Cryptanthoid genera and subgenera (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular phylogeny. Phytotaxa 318: 1 - 88. https: // doi. org / 10.11646 / phytotaxa. 318.1.1","Tabarelli, M. & Santos, A. M. M. (2004) Uma breve descricao sobre a historia natural dos brejos nordestinos, in Porto, K. C., Cabral, J. J. P. & Tabarelli, M (Eds.) Brejos de altitude em Pernambuco e Paraiba: historia natural, ecologia e conservacao. MMA / UFPE, Brasilia, pp. 17 - 24.","Silva, J. M. C. & Casteleti, C. H. M. (2005) Estado da biodiversidade da Mata Atlantica brasileira. Pp. 43 - 59 in Galindo-Leal, C. & Camara, I. G. (Eds.) Mata Atlantica, biodiversidade, ameacas e perpectivas. Fundacao SOS Mata Atlantica, Conservacao International & Centro de Ciencias Aplicadas a Biodiversidade, Belo Horizonte.","Ricketts, T. H, Dinerstein, E., Boucher, T., Brooks, T. M., Butchart, S. H. M., Hoffmann, M., Lamoreux, J. F., Morrison, J., Parr, M., Pilgrim, J. D., Rodrigues, A. S. L., Sechrest, W., Wallace, G. E., Berlin, K., Bielby, J., Burgess, N. D., Church, D. R., Cox, N., Knox, D., Loucks, C., Luck, G. W., Master, L. L., Moore, R., Naidoo, R., Ridgely, R., Schatz, G. E., Shire, G., Strand, H., Wettengel, W. & Wikramanayake, E. (2005) Pinpointing and preventing imminent extinctions. Proceedings of the National Academy of Sciences 102: 18497 - 18501. https: // doi. org / 10.1073 / pnas. 0509060102","Leme, E. M. C. & Siqueira-Filho, J. A. (2007) Atlantic Forest - an overview. In: Siqueira-Filho, J. A. & Leme, E. M. C. (Eds.) Fragments of the Atlantic Forest of Northeast Brazil, biodiversity, conservation and the bromeliads. Andrea Jakobsson Estudio, Rio de Janeiro, pp. 47 - 79.","Leme, E. M. C. & Siqueira-Filho, J. A. (2006) Taxonomia das bromelias dos fragmentos de Mata Atlantica de Pernambuco e Alagoas. In: Siqueira-Filho, J. A. & Leme, E. M. C. (Eds.) Fragmentos de Mata Atlantica do nordeste, biodiversidade, conservacao e suas bromelias. Andrea Jakobsson Estudio, Rio de Janeiro, pp. 190 - 381.","Pontes, A. R. M., Beltrao, A. C. M., Normande, I. C., Malta, A. de J. R., Silva Junior, A. P. & Santos, A. M. M. (2016) Mass extinction and the disappearance of unknown mammal species: scenario and perspectives of a biodiversity hotspot's hotspot. PloS ONE 11: e 0150887. https: // doi. org / 10.1371 / journal. pone. 0150887","Ferreira, D. M. C. & Louzada, R. B. (2020) A new species of Cryptanthus (Bromeliaceae, Bromelioideae) in the Brazilian Atlantic Forest north of the Sao Francisco River, and its contribution to the taxonomy of the genus. Systematic Botany 45: 460 - 465. https: // doi. org / 10.1600 / 036364420 X 15935294613491","Visiani, R. (1847) Semina rariora in Horto Patavino collecta. Giovanii Cecchini Tip. Venezia, pp. 4.","Visiani, R. (1854) Di Due Piante Nuove Dell'ordine Della Bromeliacee. Memorie Dell'I. R. Istituto Veneto di Cienze, Lettere ed Arti 5: 337 - 343.","Ferreira, D. M. C. (2016) Delimitacao de especies e filogeografia do complexo Cryptanthus zonatus (Vis.) Vis. (Bromeliaceae). Master thesis, Universidade Federal de Recife, Pernambuco.","Ferreira, D. M. C., Palma-Silva, C., Neri, J., Medeiros, M. C. M. P., Pinange, D. S., Benko-Iseppon, A. M. & Louzada, R. B. (2021) Population genetic structure and species delimitation in the Cryptanthus zonatus complex (Bromeliaceae). Botanical Journal of the Linnean Society 196: 123 - 140. https: // doi. org / 10.1093 / botlinnean / boaa 094","Gardner, C. S. (1986) Preliminary classification of Tillandsia based on floral characteristics. Selbyana 9: 130 - 146.","Leme, E. M. C., Halbritter, H. & Barfuss, M. H. J. (2017 a) Waltillia, a new monotypic genus in Tillandsioideae (Bromeliaceae) arises from a rediscovered, allegedly extinct species from Brazil. Phytotaxa 299: 1 - 35. https: // doi. org / 10.11646 / phytotaxa. 299.1.1","Lindley, J. (1843) Vriesea psittacina, Parrot-flowered Vriesea. Edward's Botanical Register 29: 9 - 11.","Harms, H. (1929 b) Bromeliaceae novae III. Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 10: 784 - 805. https: // doi. org / 10.2307 / 3994800","Carvalho, J. D. T., Oliveira, J. M. S. de, Freitas, C. C. de & Martins, M. S. (2016) Stamen morphoanatomy of Dyckia Schult. F. (Bromeliaceae, Pitcairnioideae) species: new data for taxonomic use. Acta Botanica Brasilica 30: 389 - 400. https: // doi. org / 10.1590 / 0102 - 33062016 abb 0112","Carvalho, J. D. T., Essi, L. & Oliveira, J. M. S. (2017) Flower and floral trichome morphology of species of Dyckia Schult. f. (Bromeliaceae, Pitcairnioideae), and their importance to species characterization and genus taxonomy. Acta Botanica Brasilica 31: 29 - 41. https: // doi. org / 10.1590 / 0102 - 33062016 abb 0335","Buneker, H. M. (2021) Revisao taxonomica do complexo Dyckia selloa (Bromeliaceae, Pitcairnioideae) baseada em caracteres morfologicos. Master's thesis, Universidade Federal do Rio Grande do Sul, Porto Alegre, 231 pp.","Leme, E. M. C, Zizka, G., Paule, J., Aguirre-Santoro, J., Heller, S., Ramirez-Morillo, I. M., Halbritter, H., Mariath, J. E. A., Carvalho, J. D. T & Forzza, R. C. (2021) Re-evaluation of the Amazonian Hylaeaicum (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular evidence. Phytotaxa 499: 1 - 60. https: // doi. org / 10.11646 / phytotaxa. 499.1.1","Mez, C. (1934 - 1935) Bromeliaceae. In: Engler, A. (Ed.) Das Pflanzenreich, regni vegetabilis conspectus 4 (32). W. Engelmann, Leipzig, 667 pp.","Barfuss, M. H. J., Till, W., Leme, E. M. C., Pinzon, J. P., Manzanares, J. M., Halbritter, H., Samuel, R. & Brown, G. K. (2016) Taxonomic revision of Bromeliaceae subfam. Tillandsioideae based on a multi-locus DNA sequence phylogeny and morphology. Phytotax a 279: 1 - 97. https: // doi. org / 10.11646 / phytotaxa. 279.1.1","Varadarajan, G. S. & Brown, G. K. (1988) Morphological variation of some floral features of the subfamily Pitcairnioideae (Bromeliaceae) and their significance in pollination biology. Botanical Gazette 149: 82 - 91. https: // doi. org / 10.1086 / 337694","Brown, G. K. & Gilmartin, A. J. (1989) Stigma types in Bromeliaceae: a systematic survey. Systematic Botany 14: 110 - 132. https: // doi. org / 10.2307 / 2419055","Leme, E. M. C. (1997) Canistrum - Bromeliads of the Atlantic Forest, Salamandra, Rio de Janeiro, pp. 1 - 107.","Leme, E. M. C. (1998) Canistropsis - Bromeliads of the Atlantic Forest, Salamandra, Rio de Janeiro, pp. 1 - 143.","Leme, E. M. C. (2000) Nidularium - Bromeliads of the Atlantic Forest, Sextante, Rio de Janeiro, pp. 1 - 263.","Turland, N. J., Wiersema, J. H., Barrie, F. R., Greuter, W., Hawksworth, D. L., Herendeen, P. S., Knapp, S., Kusber, W. - H., Li, D. - Z., Marhold, K., May, T. W., McNeill, J., Monro, A. M., Prado, J., Price, M. J. & Smith, G. F. (eds.) (2018) International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashutten: Koeltz Botanical Books. https: // doi. org / 10.12705 / Code. 2018"]}

Orthocryptanthus santaritensis (Leme, S. Heller & Zizka) Leme, Zizka & Paule, comb. nov. Basionym:— Orthophytum santaritense Leme, S. Heller & Zizka, Phytotaxa 318: 78. 2017. Type:— BRAZIL. Minas Gerais: Santa Rita do Itueto, Pedra de Santa Rita do Itueto, ca. 1020 m elevation, 19º 22’ 24.64” S, 41º 21’ 30.49” W, 23 September 2014, R. Vasconcelos Leitão & L. C . Medeiros s.n., fl. cult. September 2014, E . Leme 8945 (holotype RB!, isotype HB!)., Published as part of 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, pp. 128-170 in Phytotaxa 544 (2) on page 148, DOI: 10.11646/phytotaxa.544.2.2, http://zenodo.org/record/6503832

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., Published as part of 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, pp. 128-170 in Phytotaxa 544 (2) on pages 145-148, DOI: 10.11646/phytotaxa.544.2.2, http://zenodo.org/record/6503832, {"references":["Leme, E. M. C. Heller, S., Zizka, G. & Halbritter, H. (2017 b) New circumscription of Cryptanthus and new Cryptanthoid genera and subgenera (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular phylogeny. Phytotaxa 318: 1 - 88. https: // doi. org / 10.11646 / phytotaxa. 318.1.1","Schulte, K. & Zizka, G. (2008) Multi locus plastid phylogeny of Bromelioideae (Bromeliaceae) and the taxonomic utility of petal appendages and pollen characters. Candollea 63: 209 - 225.","Aguirre-Santoro, J., Betancur, J., Brown, G. K., Evans, T. M., Salgueiro, F., Alves-Ferreira, M. & Wendt, T. (2015) Is Ronnbergia (Bromeliaceae, Bromelioideae) a geographically disjunct genus? Evidence from morphology and chloroplast DNA sequence data. Phytotaxa 219: 261 - 275. https: // doi. org / 10.11646 / phytotaxa. 219.3.6","Aguirre-Santoro, J., Michelangeli, F. A. & Stevenson, D. W. (2016) Molecular phylogenetics of Ronnbergia alliance (Bromeliaceae, Bromelioideae) and insights into their morphological evolution. Molecular Phylogenetics and Evolution 100: 1 - 20. https: // doi. org / 10.1016 / j. ympev. 2016.04.007","Morren, E. & Andre (1874) Ronnbergia morreniana Linden & Andre, Ronnbergia de Morren, Bromeliacees. In: Linden, J. J. & Andre, E. (Eds.) L'Illustration Horticole 21: 120.","Sass, C. & Specht, C. D. (2010) Phylogenetic estimation of the core Bromelioids with an emphasis on the genus Aechmea (Bromeliaceae). Molecular Phylogenetics and Evolution 55: 559 - 571. https: // doi. org / 10.1016 / j. ympev. 2010.01.005","Leme, E. M. C, Ribeiro, O. B. C, Souza, F. V. D., Souza, E. H., Kollmann, L. J. C. & Fontana, A. P. (2020) Miscellaneous new species in the \" Cryptanthoid complex \" (Bromeliaceae: Bromelioideae) from Eastern Brasil. Phytotaxa 430: 157 - 202. https: // doi. org / 10.11646 / phytotaxa. 430.3.2"]}

Orthocryptanthus vasconcelosianus (Leme) Leme, Zizka & Paule, comb. nov. Basionym:— Orthophytum vasconcelosianum Leme, Phytotaxa 205: 287. 2015. Type:— BRAZIL. Minas Gerais: Alvarenga, Córrego Sujo, hill at the foot of Pico da Aliança, ca. 1060 m elevation, 19º 23’ 21” S, 41º 41’ 10” W, 23 June 2012, E. Leme 8673, R . Vasconcelos Leitão & R . Faria (holotype RB!, isotype HB!)., Published as part of 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, pp. 128-170 in Phytotaxa 544 (2) on page 149, DOI: 10.11646/phytotaxa.544.2.2, http://zenodo.org/record/6503832

Siqueiranthus cinereus (D.M.C. Ferreira & Louzada) Leme, Zizka, E.H. Souza & Paule, comb. nov. Basionym:— Cryptanthus cinereus D.M.C. Ferreira & Louzada, Systematic Botany 45: 460. 2020. Type:— BRAZIL. Alagoas: Ibateguara, Sítio Duas Barras, 8 May 2019, D. Cavalcanti, E. M. de Almeida, A. Melo, F. G. Silva, W. T. C. C. Santos & V. B. V. Souza 921 (holotype: UFP!, isotypes: MAC, RB, UFRN, SP, NY, unavailable). Description:— Plants andromonoecious, saxicolous, long caulescent usually by the successive growth of a single apical shoot, stems 18–160 cm long, 1.7–2.8 cm in diameter, rigid, erect to prostrate, propagation by a single shoot at the base of the inflorescence and by few shoots along the stem, rhizomes not detected. Leaves 30–49 in number, equally arranged along the stem, strongly coriaceous; sheath suborbicular to reniform-trapeziform, 2.8–3.5 × 4.4– 6 cm, yellowish and glabrous near the base, green or reddish and densely cinereous lepidote toward the base and mainly abaxially strongly corrugate, persisting on the older part of the stem, densely spinose distally; spines narrowly triangular, 0.5–3 mm long, 0.2–3 mm apart, straight, antrorse or retrorse, densely white lepidote; blade 17–57 × 4–4.7 cm, narrowly triangular-lanceolate, attenuate-acuminate, suberect to spreading or reflexed (mainly at anthesis), green or reddish-castaneous when fully exposed to sun, abaxial surface completely covered by a dense layer of cinereous trichomes obscuring the blade color, adaxial surface densely lepidote by cinereous trichomes obscuring to not at all obscuring the blade color, becoming partially glabrescent with age, margins straight to slightly undulate, densely spinose near the base to laxly spinulose toward the apex; spines triangular, 0.5–3 mm long, 2–14 mm apart, antrorse. Inflorescence compound, sessile, corymbose or a compact glomerulate spike, 4–6 cm long (petals not included), 4–4.5 cm in diameter, with perfect and staminate flowers in the lateral fascicles, bearing a distinct apical-central simple head of 19–25 staminate flowers; primary bracts foliaceous or subfoliaceous, the basal ones many times exceeding the fascicles, the upper ones abruptly reduced in size, slightly exceeding the fascicles, narrowly triangular, green or becoming rose to vinaceous at anthesis, forming an inconspicuous colorful ring around the inflorescence in contrast with the prevailingly cinereous-green color of its distal portion, both surfaces densely cinereous lepidote, margins straight to undulate, densely spinose near the base to laxly spinulose toward the apex; spine s 0.5–3 mm long, 1–10 mm apart, antrorse; fascicles 5–7 in number, with 2–7 perfect and/or staminate flowers, complanate, flabellate, 40–45 × 20–25 mm (petals excluded), subsessile to shortly stipitate, with 2–4 lateral sterile bracts; stipe subcomplanate, 5–6 × 4–5 mm. Floral bracts sublinear to narrowly triangular-lanceolate, cymbiform, densely and coarsely white lepidote toward the apex, hyaline to greenish-white near the base and green to rose toward the apex, margins entire or inconspicuously and sparsely denticulate, those in the fascicles 20–34 × 5–13 mm, strongly carinate to ecarinate, apex acute, those in the staminate flowers of the apical/simple part of the inflorescence 20–27 × 5–7 mm, ecarinate to obtusely carinate, apex acuminate. Flowers sessile, diurnal, fragrance not detected, the perfect ones 45–52 mm long (including the petals), the staminate ones 30–48 mm long (including the petals); sepals narrowly lanceolate, acuminate, thin in texture, carinate to obtusely carinate, whitish near the base, green to rose toward the apex, densely and coarsely white lepidote with brown-centered trichomes, obscuring the sepal color at the apex, entire, those of the perfect flowers 21–22.2 × 3–5 mm, connate at the base for (2.4-) 5–6 mm, those of the staminate flowers 16–22.2 × 3–5 mm, connate at the base for 4–5 mm; petals narrowly obovate to oblanceolate, rounded to slightly emarginate, free or connate for less the 1 mm (subfree), erect except for the suberect apex, forming a prevailingly tubular corolla, 20–26 × 4–6 mm, bearing well developed appendages as well as 2 conspicuous longitudinal callosities about reaching the midpoint of the filaments; appendages scutelliform with tendency to cupuliform, thick, 3–5 mm above the base, upper and lower margin inconspicuously and irregularly scalloped, not glandulose; stamens included, partially visible inside the corolla at anthesis; filament white, terete, unequal, 11–18 mm long, the antesepalous ones free, longer than the antepetalous ones, the antepetalous ones adnate to the petals for 1–3 mm; anther 2.5–5 mm long, base deeply bilobed, apex obtuse and inconspicuously apiculate, fixed at the base of the connective between the basal lobes (basifixed), not laterally compressed at anthesis, dehiscence introrse; pollen sulcate, spherical, ca. 55 µm, reticulate, reticulum with slightly flat muri and broad lumina decreasing in size towards the sulcus margins especially towards the apices, sulcus broad, without distinct margins, covered with exine elements partially forming a net. Style 20–25.5 mm long, completely reduced in straminate flowers; stigma simple-erect with tendency to simple-patent, white, lobes longer than broad, elliptic or narrowly obovate in outline, suberect, 2–2.2 × 1 mm, navicular, V-shaped with a central furrow, bearing regular and fine transversal grooves from the middle toward the margins and with a slightly undulate surface, margins inconspicuously crenulate, with sparse and irregular long filamentous white trichomes. Ovary trigonous, white, glabrous to inconspicuously and sparsely white lepidote with fimbriate trichomes, in the perfect flowers 15–19 × 6–8.2 mm, ovoid, in the staminate flowers 10–16 × 4–6 mm, cylindrical; epigynous tube 5–8 mm, narrow; ovules obtuse, 18–25 per locule in perfect flowers, fewer in number to absent in the staminate flowers; placentation apical. Fruits 18–25 × 7–8 (–9.2) mm, narrowly obovate to ellipsoid, trigonous, glabrous to inconspicuously white lepidote, white except the greenish or reddish-green distal portion, sepals in fruit persistent, stramineous, the distal portion often decaying, remnants 6–7 times shorter than the fruit length; seeds 10–20 per fruit, 2–3.5 (–4) × 1.5–3 mm, asymmetrical, subquadrate, subreniform or subtrapeziform, pale brown when mature, distinctly striate. Distribution and habitat:— Siqueiranthus cinereus is only known from the type population, as a saxicole, growing at the top of a granitic outcrop in the Atlantic Forest domain in the municipality of Ibateguara, Alagoas state, Brazil (fig. 11.). The area is part of a small but rich fragment of vegetation located inside farms and isolated by surrounding pasturelands (fig. 11 F). It is accessible to horses, cows, and other domestic animals, which were seen eating leaves and inflorescences of S. cinereus, certainly causing an important impact on the reproduction and survival of its tiny population. Probably, this fragment will be destroyed in the coming years and completely occupied by pasturelands, despite its rich bromeliad community composed by endemic and even endangered species, like Aechmea catendensis Siqueira & Leme (Leme & Siqueira-Filho, 2006: 205), A. costantinii (Mez, 1916: 245) Smith (1970: 281), A. leptantha (Harms, 1929a: 786) Leme & Siqueira (2006: 213), A. nudicaulis var. nordestina Siqueira & Leme (Leme & Siqueira, 2006: 243), besides several tillandsioid species, to name a few. The investigation performed in the ecologically similar hills around the type locality did not reveal any subpopulation of S. cinereus (J.A. Siqueira pers. comm.). The studied population at the type locality ‒ the only known one ‒ is limited in the number of plants, being composed by approximately 800 individuals in different stage of development, most of them wih mature size. Seedlings were not observed at the spot. The most part of the population is partially or completely shaded by shrubs and short trees of Melastomataceae, Clusiaceae, Myrtaceae, Rubiaceae, and Asteraceae, being intermingled with them. Without a strong enough root system to support the upright growth of the long-caulescent plants, S. cinereus grow propped up by the surrounding bushes (fig. 11 D). At the periphery of the vegetation island, the leaves of some individuals of S. cinereus, exposed to full sun, turn reddish. Plotting the known occurrence of S. cinereus in Geocat software (Bachman et al. 2011), the obtained estimated extent of occurrence (EOO) is 0.051 km ², and the area of occupancy (AOO) is 8 km ², placing the species in the IUCN concept of being a critically endangered (CR) species. When its small area of occurrence is associated to the highly negative human pressure mentioned above, that affects S. cinereus, according to the criteria B1 [ab (i, ii, iii)] of the IUCN (2020) its critically endangered (CR) status is reiterated. So, the in-situ survival of this unusual species is uncertain, recommending extra conservation efforts to keep it alive at least ex-situ. Etymology:—The epithet of this species name is a clear reference to the dense cinereous coverage of trichomes that characterizes it. Distinctive characters:— Siqueiranthus cinereus is a unique species without close relationship with any known species of the “Cryptanthoid complex” in general or specifically from the region north of São Francisco river where it lives. Its uniqueness was highlighted in the protologue concerning leaf blade width and dense layer of cinereous trichomes (Ferreira & Louzada 2020), and it was then associated to the endemic Cryptanthus felixii Siqueira & Leme (Leme & Siqueira 2006: 285), which is morphologically and phylogenetically not related to it. Differences to Cryptanthus are indicated in the respective item above., Published as part of 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, pp. 128-170 in Phytotaxa 544 (2) on pages 159-160, DOI: 10.11646/phytotaxa.544.2.2, http://zenodo.org/record/6503832, {"references":["Leme, E. M. C. & Siqueira-Filho, J. A. (2006) Taxonomia das bromelias dos fragmentos de Mata Atlantica de Pernambuco e Alagoas. In: Siqueira-Filho, J. A. & Leme, E. M. C. (Eds.) Fragmentos de Mata Atlantica do nordeste, biodiversidade, conservacao e suas bromelias. Andrea Jakobsson Estudio, Rio de Janeiro, pp. 190 - 381.","Mez, C. (1916) Additamenta monographica 1916. Repertorium Specierum Novarum Regni Vegetabilis 14: 241 - 256. https: // doi. org / 10.1002 / fedr. 19160141603","Smith, L. B. (1970) Notes on Bromeliaceae, XXX. Phytologia 19: 281 - 290. https: // doi. org / 10.5962 / bhl. part. 14773","Harms, H. (1929 a) Bromeliaceae novae II. Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 10: 573 - 582. https: // doi. org / 10.2307 / 3994782","Bachman, S., Moat, J., Hill, A. W., Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. In: Smith, V. & Penev, L. (Eds.) e-Infrastructures for data publishing in biodiversity science. ZooKeys 150: 117 - 126. https: // doi. org / 10.3897 / zookeys. 150.2109","IUCN (2020) The IUCN Red List of Threatened Species. Version 2020 [online]. Available from: http: // www. iucnredlist. org (Accessed 4 October 2021).","Ferreira, D. M. C. & Louzada, R. B. (2020) A new species of Cryptanthus (Bromeliaceae, Bromelioideae) in the Brazilian Atlantic Forest north of the Sao Francisco River, and its contribution to the taxonomy of the genus. Systematic Botany 45: 460 - 465. https: // doi. org / 10.1600 / 036364420 X 15935294613491"]}

Orthophytum caraibense Leme & O.B.C. Ribeiro, sp. nov. (figs. 19 A to H) Diagnosis:—This new species is morphologically related to O. duartei, but differs by its shorter stature (7–8 cm vs. 18–36 cm tall), propagation by short basal shoots (vs. slender stolons), leaf blades smaller (10–14 × 1–1.8 cm vs. 15–25 × 2–3 cm), shorter peduncle (1–1.5 cm vs. 5–7 cm long), shorter inflorescence (4–6 cm vs. 12–25 cm long), longer flowers (40–43 mm vs. 26–27 mm long), petals longer (35–39 mm vs. × 20–21 mm long), exappendiculate (vs. bearing distinct cupuliform appendages), apex acuminate (vs. rounded), and stigma conduplicate-patent (vs. simple-patent). Type:— Brazil. Minas Gerais: Bonito de Minas, RPPN Vereda da Caraíba, saxicolous on sandstone outcrops in Cerrado vegetation, 780 m elevation, 14°43’11.79” S, 45°02’03.87” W, 27 January 2019, C . V. Vidal s.n., cult. E . Leme 9534 (holotype RB!). Description:— Plants stemless, 7–8 cm long when flowering, propagating by short basal shoots. Leaves 8–10 in number, not distinguishable from the primary bracts at anthesis, arcuate to subspreading at anthesis, forming a narrow rosette; sheath inconspicuous; blade narrowly lanceolate, apex caudate, 10–14 × 1–1.8 cm, thin coriaceous, canaliculate, green to greenish-bronze toward the apex, abaxially densely to subdensely and coarsely white lepidote, nerved, adaxially glabrous, margins straight to slightly undulate, densely to subdensely spinose; spines acicular, prevailingly straight to slightly antrorsely uncinate, 1–2 mm long, 2–5 mm apart. Inflorescence compound, 4–6 cm long (excluding the petals), 2–2.5 cm in diameter, slightly to distinctly elongated; peduncle inconspicuous, 1–1.5 × 0.7 cm, white lepidote; peduncle bracts foliaceous, densely arranged, arcuate to subspreading; primary bracts foliaceous, green, distinctly exceeding the fascicles, suberect to spreading at anthesis; fascicles sessile, inconspicuous, subflabellate, 5–7 in number, 2–4-flowered. Floral bracts narrowly triangular, attenuate-caudate, slightly to distinctly shorter than the sepals, 20–24 × 5–7 mm, thin in texture, greenish-white, glabrous or centrally with sparse and inconspicuous glandulose trichomes, carinate, nerved, entire. Flowers sessile, 40–43 mm long (with extended petals), slightly fragrant, anthesis starting at night and continuing during the next day; sepals subsymmetric, narrowly lanceolate, attenuate-caudate, erect, 21–27 × 2–3.5 mm, free, greenish-white or reddish toward the apex, glabrous or sometimes with inconspicuous and sparse fimbriate white trichomes at the apex, entire, ecarinate, membranaceous; petals narrowed toward the base, with a distinct narrowly ovate-lanceolate blade, apex acuminate, 35–39 × 5–5.5 mm, free, white, spreading at anthesis and forming a fan blade-like corolla of 20–25 mm in diameter, exappendiculate, bearing 2 longitudinal callosities about equaling the filaments and thickened from about 5 mm above the base. Stamens included but partially visible at anthesis; filament distinctly unequal in length, the antesepalous ones 24–27 mm long, free, the antepetalous ones 20–22 mm long, ca. 0.3 mm in diameter, adnate to the petals for 15–17 mm; anther 1.2–2 mm long, dorsifixed near the base, base obtuse, apex obtuse and inconspicuously apiculate, strongly laterally compressed at anthesis. Ovary ca . 4.5 × 3 mm, white, glabrous, trigonous; epigynous tube inconspicuous, ca. 0.5 mm long; stigma conduplicate-patent, lobes narrow, spreading-recurved, ca. 0.4 mm long, white, distinctly exceeding the anthers and visible above the corolla throat, margins irregularly and distinctly scalloped, undulate, inconspicuously and sparsely papillate toward the distal end; ovules many, obtuse; placentation subapical. Fruits baccate, 5–6.5 × 3–5 mm, white, with persistent sepals, 3–4.5 times longer than the fruit length; seeds 1–1.5 × 0.7–1 mm, 50–56 in number, subtriangular, proximal end acute, distal end obtuse. Distribution and habitat:— Orthophytum caraibense grows in the north of Minas Gerais state, close to the border with Bahia state, with a geographical range extending much inland when compared to the closest morphological relative, O. duartei, which lives in the Atlantic Forest domain of Espírito Santo state, closer to the Atlantic coast. This new species lives as a saxicole in crevices in vertical sandstone outcrops in Cerrado vegetation (figs 14 A–C), at an altitude of ca. 780 m. The size of the population and the extension of the occupied area is not clearly known, and so it is considered here a “data deficient” and its conservation status is inconclusive, despite its occurrence inside a private protected area. Etymology:—The name chosen for this new species is a reference to the private reserve ‒ Reserva Particular do Patrimônio Natural (RPPN) Vereda da Caraíba ‒ where this new species was discovered. Distinctive characters:— Orthophytum caraibense is an outlier species, like its closest morphological relative O. duartei, mainly due to its unappendaged petals (fig. 19 G) and conduplicate-patent stigma type (fig. 19 H) not observed in any other member of Orthophytum. Exappendiculate petals are also observed in Cryptanthus, Forzzaea, Hoplocryptanthus, and frequently in Orthocryptanthus, while the conduplicate-patent stigma type is only seen in Cryptanthus and Lapanthus, Krenakanthus has the conduplicate-spiral type. However, all other floral features are in accordance with Orthophytum. According to the molecular phylogeny, both O. caraibense and O. duartei are part of an unsupported clade (0.53 PP, 57 BS) along with Orthophytum subgg. Orthophytum, Capixabanthus and Clavanthus (fig. 1). Orthophytum duartei remains without or with weak support (0.73 PP, 67 BS) as sister to Orthophytum subg. Capixabanthus and both are unsupported sister to the Orthophytum-Clavanthus clade. However, Orthophytum subgg. Orthophytum and Clavanthus are sister to each other with high support (1 PP, 98 BS). The position of O. caraibensis in the phylogenetic tree is challenging. Despite of being well supported (1 PP, 98 BS), in all aspects its geographical range and morphology are not compatible with subgenus Clavanthus, to which it comes out in sister position. Apart of its much smaller size (7–8 cm vs. 18–36 cm tall) and leaf blades (10–14 × 1–1.8 cm vs. 15–25 × 2–3 cm), the thin coriaceous leaf texture and the fan blade-like corolla with spreading petals of O. caraibense are similar to O. duartei (see Leme 2014, Leme et al. 2017b). This new species also differs from its closest relatives in the propagation by short basal shoots (vs. slender stolons), shorter peduncle (1–1.5 cm vs. 5–7 cm long), shorter inflorescence (4–6 cm vs. 12–25 cm long), longer flowers (40–43 mm vs. 26–27 mm long), longer sepals (21–27 mm vs. 10–12 mm), longer (35–39 mm vs. 20–21 mm long) and exappendiculate petals (vs. bearing distinct cupuliform appendages), with apex acuminate (vs. rounded), and stigma of the conduplicate-patent type (vs. simple-patent). The reported affinity of O. caraibense and O. duartei and their equivocal position in the molecular phylogeny requires further investigation and refinement of the molecular approach and for the time being we therefore refrain from elevating Capixabanthus and Clavanthus to generic status, despite its well-circumscribed morphology and welldelimited biogeography., Published as part of 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, pp. 128-170 in Phytotaxa 544 (2) on pages 160-165, DOI: 10.11646/phytotaxa.544.2.2, http://zenodo.org/record/6503832, {"references":["Leme, E. M. C. (2014) Studies on Orthophytum - Part XII: rediscovering Orthophytum duartei. Journal of the Bromeliad Society 64: 176 - 184.","Leme, E. M. C. Heller, S., Zizka, G. & Halbritter, H. (2017 b) New circumscription of Cryptanthus and new Cryptanthoid genera and subgenera (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular phylogeny. Phytotaxa 318: 1 - 88. https: // doi. org / 10.11646 / phytotaxa. 318.1.1"]}

Krenakanthus roseolilacinus (Leme) Leme, Zizka & Paule, comb. nov. Basionym:— Orthophytum roseolilacinum Leme, Phytotaxa 205: 284. 2015. Type:— BRAZIL. Minas Gerais: Conselheiro Pena, Serra do Padre Ângelo, Mata do Elias, 810 m elevation, 19° 17’ 14.71” S, 41° 34’ 13.47” W, May 2014, R. Vasconcelos Leitão & E.C. Ribeiro s.n., cult. E . Leme 8885 (holotype RB!). Description:— Plants terrestrial, monoecious, long caulescent, flowering 30–60 cm tall, propagating by elongated shoots developing near the base of the inflorescence. Leaves 20–27 in number, thin in texture, chartaceous, except for a thicker median zone mainly toward the base, spreading-recurved, more or less equally arranged along the stem, forming a subdense rosette; sheath trapeziform, ca. 2 × 3.5 cm, pale green, glabrescent, densely spinulose; blade narrowly lanceolate, long attenuate then caudate, not narrowed at the base, 17–27 × 1.8–2.4 cm, green, concolorous, glabrescent to glabrous, margins undulate mainly toward the base, densely spinulose; spines pale green, the basal ones narrowly triangular, 1–3 mm long, 2–4 mm apart, spreading to slightly antrorse, the upper ones triangular, 0.2–0.3 mm long, 1–3 mm apart, antrorse. Inflorescence sessile, 3.5–5 cm long, corymbose, sparsely branched only at the base and simple toward the apex; primary bracts foliaceous, not contrasting in color with the leaf blades spreading-recurved, many times exceeding the fascicles; flower fascicles only 1–2 in number, densely aggregated, 30–32 × 14–16 mm (excluding the petals), ca. 7 mm thick, 2–4-flowered, stipes inconspicuous, stout, ca. 5 mm long; floral bracts in the fascicles triangular, acuminate-caudate, membranaceous, green toward the apex, hyaline near the base, glabrous, nerved, slightly shorter than the sepals, alate-carinate (the outer ones) to obtusely if at all carinate (the inner ones), 20– 26 × 14–15 mm, margins entire to spinulose at the apex; floral bracts in the unbranched apical part of the inflorescence resembling the leaves, many times exceeding (basal ones) to slightly shorter than the flowers (apical ones). Flowers all perfect, sessile, 42–55 mm long, fragrant; sepals 20–27 × 4–5 mm, sublinear-lanceolate to narrowly ovate-lanceolate, acuminate-caudate, connate at the base for 3–7 mm, green but drying dark castaneous soon after anthesis, glabrous, thin in texture, the adaxial ones alate-carinate with the keel decurrent on the ovary, the abaxial one obtusely if at all carinate; petals broadly spathulate from a very narrow base, 35–44 × 15–20 mm, 1.8–2.9 times longer than wide, free, greenish-white toward the base, rose-lilac to lilac-purple toward the apex, the blades suborbicular, broadly acute to rounded, spreading at anthesis and flaccidescent afterwards, the distal margins bearing sparse and inconspicuous glandular trichomes, bearing 2 longitudinal callosities at the base, equaling to exceeding the antesepalous filaments, and 2 appendages at the base; appendages ca . 2.5 mm above the base, thick, cupuliform, margins downwardly curved, minutely and obtusely crenulate. Stamens deeply included and not visible; filament distinctly unequal in length, the antepetalous ones 4–6 mm long, adnate to the petals for ca. 1.5 mm, the antesepalous ones 11–14 mm long, free; anther 1.5–2 mm long, oblong, dorsifixed at ca. 1/3 from the base, base bilobed, apex obtuse or minutely apiculate; pollen sulcate, oblate, ca. 45 μm long, reticulate, reticulum broadly meshed proximally and meshes distinctly decreasing in width towards the sulcus margins, the sulcus narrow, covered by small exine elements, margins weakly defined; style slightly exceeding the antepetalous stamens and distinctly shorter than the antesepalous ones, white; stigma conduplicate-spiral, lobes elongated, loosely contorted, densely papillose; ovary 5–7 mm long, 5–8 mm in diameter at the apex, trigonous, white, glabrous; epigynous tube lacking; ovules numerous, obtuse; placentation median to apical. Fruits baccate, subglobose, whitish toward the base, yellowish toward the apex, obtusely angulose, 8–10 × 7–12 mm, sepals persistent, 2.5–3.5 times longer than the fruit length; seeds few to numerous, narrowly ovoid to fusiform, 2–3 × 0.7–1 mm, slightly to distinctly curved, distinctly sulcate, dark brown, bearing a unilateral, thick, greenish-transluscent raphe., Published as part of 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, pp. 128-170 in Phytotaxa 544 (2) on page 145, DOI: 10.11646/phytotaxa.544.2.2, http://zenodo.org/record/6503832

Orthocryptanthus arcanus (Leme) Leme, Zizka & Paule, comb. nov. Basionym:— Orthophytum arcanum Leme, Phytotaxa 430: 191. 2020. Type: — BRAZIL. Minas Gerais: Conselheiro Pena, near the top of the Pico do Padre Angelo, 1441 m elevation, 19º18’59” S, 41º34’44” W, September 2015, R. Vasconcelos Leitão s.n., fl. cult. E . Leme 9093 (holotype RB!)., Published as part of 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, pp. 128-170 in Phytotaxa 544 (2) on page 148, DOI: 10.11646/phytotaxa.544.2.2, http://zenodo.org/record/6503832

Key to Cryptanthoid genera and subgenera 1 Plants andromonoecious with perfect flowers in the basal fascicles and the staminate ones concentrated in the central/apical portion of the inflorescence; pollen sulcus partially or totally covered by exine elements forming a net; fruits 12–25 mm long, with the distal portion of the persistent sepals usually soon decaying....................................................................................................... 2 1* Plants with perfect flowers only; pollen sulcus only covered by small and sparse exine elements, never forming a net; fruits 4–10 mm long, with persistent sepals.......................................................................................................................................................... 3 2 Petals distinctly connate, unappendaged, arcuate-recurved at anthesis and forming a fan blade-like corolla; filaments equal; stigma conduplicate-patent; fruits 12–20 × 9–12 mm; epigynous tube usually lacking or sometimes insconspicuous (to 3 mm long).................................................................................................................................................................................................. Cryptanthus 2* Petals free or subfree, appendaged, erect except for the suberect apex, forming a prevailingly tubular corolla; filaments unequal; stigma simple-erect with tendency to simple-patent; fruits 18–25 × 7–8 (–9.2) mm; epigynous tube 5–8 mm long.. Siqueiranthus 3 Petals without appendages, sometimes with well developed longitudinal callosities........................................................................ 4 3* Petals with well developed appendages and sometimes with longitudinal callosities as well........................................................... 8 4 Petals usually white, connate at the base or rarely free but then the petals greenish-yellow in their visible parts............................ 5 4* Petals free, white or lilac-rose............................................................................................................................................................ 7 5 Basal flower fascicles with (5-) 6–15 flowers; pollen 55–60 µm; stigma simple-imbricate; plants from the Atlantic Forest of Espírito Santo.................................................................................................................................................................. Rokautskyia 5* Basal flower fascicles with 2–5 flowers or the inflorescence simple; pollen 40–50 µm; stigma not simple-imbricate; plants from Campos Rupestres of Minas Gerais................................................................................................................................................... 6 6 Leaves coriaceous, not succulent; petals connate at the base to 1/3 of their length, or rarely free but then the petals greenish-yellow in the visible parts; anthers always straight at anthesis; pollen ca. 50 µm; stigma cylindric-distent; persistent sepals slightly shorter to equaling the fruit length or rarely 1.4 times longer; seeds 35–75 per fruit........................................................ Hoplocryptanthus 6* Leaves thick-coriaceous, succulent; petals connate at the base to 1/7 of their length; anthers usually strongly recurved to spirally coiled at anthesis; pollen ca. 40 µm; stigma simple-erect or simple-patent with a tendency to simple-dilated; persistent sepals up to 1.5 times longer than the fruit; seeds 3–8 per fruit............................................................................................................ Forzzaea 7 Plants stemless; leaves thick-coriaceous, succulent; inflorescence with inconspicuous, to 4-flowered fascicles; stamens subequal in length; anthers usually strongly recurved to spirally coiled at anthesis; pollen ca. 40 µm............................................... Forzzaea 7* Plants distinctly caulescent; leaves coriaceous, not succulent; inflorescence with conspicuous, up to 9-flowered fascicles; stamens distinctly unequal with the antepetalous ones much shorter than the antesepalous ones; anthers straight; pollen 45–50 µm............................................................................................................................................................................................. Orthocryptanthus 8 Petals with laminiform appendages; stigma conduplicate-patent...................................................................................... Lapanthus 8* Petals appendages not laminiform; stigma simple-dilated, simple-patent or conduplicate-spiral..................................................... 9 9 Inflorescence sessile or inconspicuously pedunculate and petals neither obtuse-cucullate nor forming a clavate or subclavate corolla.............................................................................................................................................................................................. 12 9* Inflorescence on a short but distinct to elongate peduncle, or if sessile then the plants long caulescent, and with the basal portion of the central leaves and the primary bracts turning reddish or bright red, forming a colorful ring around the inflorescence, petals obtuse-cucullate and forming a subclavate corolla.......................................................................................................................... 10 10 Plants long caulescent and the basal portion of the central leaves and primary bracts turning reddish or bright red at anthesis, forming a colorful ring around the inflorescence, if short caulescent or stemless then the petal appendages of the cupuliform or sacciform type; pollen 35–40 µm in diameter........................................................................... Orthophytum subg. Capixabanthus 10* Plants stemless or nearly so, sometimes pseudocaulescent but then without a distinct rosette; petal appendages of the echinatiform or scutelliform type, rarely with tendency to cupuliform or sacciform; pollen 40–60 µm in diameter........................................... 11 11 Flower fascicles subflabellate-pulvinate; petals obtuse-cucullate, erect and forming a clavate corolla not exposing the stamens; petal appendages scutelliform or rarely sacciform; plants from the central-northern Espinhaço Range in Minas Gerais state.................................................................................................................................................................... Orthophytum subg. Clavanthus 11* Flower fascicles usually strobilate; petals acuminate, acute or rounded, erect except for the suberect to recurved distal portion, exposing the stamens; petal appendages echinatiform; plants with much broader geographical range, but mostly not encompassing the Espinhaço Range in Minas Gerais state.................................................................................. Orthophytum subg. Orthophytum 12 Plants long caulescent; petals broadly spathulate from a very narrow base, blades suborbicular, rose-lilac to lilac-purple, spreading at anthesis and flaccidescent afterwards; stamens deeply included and not visible; stigma conduplicate-spiral...................................................................................................................................................................................................................... Krenakanthus 12* Plants stemless; petals narrowly spathulate, blades ovate to obovate, white, erect to recurved at anthesis, not flaccidescent and remaining erect or nearly so afterwards; stamens visible at least in part; stigma simple-erect to simple-patent............................. 13 13 Leaves 10–12 in number; primary bracts sometimes red near the base at anthesis but not forming a colorful ring around the inflorescence; endemic to the mesoregion of the Rio Doce valley, Minas Gerais, in the Atlantic Forest domain.............................................................................................................................................................................................................. Orthocryptanthus 13* Leaves 25–70 in number; primary bracts and the basal portion of the inner leaves turning white, yellow or red, forming a colorful ring around the inflorescence in contrast with the color of the distal portion of the leaves; endemic to the Campos Rupestres of the Septentrional Plateau of the Espinhaço Range in Minas Gerais and Bahia states.............................................................. Sincoraea, Published as part of 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, pp. 128-170 in Phytotaxa 544 (2) on pages 165-166, DOI: 10.11646/phytotaxa.544.2.2, http://zenodo.org/record/6503832

Hylaeaicum stoloniferum (L.B. Sm.) Leme, Zizka & Aguirre-Santoro, comb. nov. Basionym:— Neoregelia stolonifera L.B. Sm., Phytologia. 9: 246, pl. 1, figs. 13, 14. 1963. Type:— PERU. Loreto: Vicinity of Iquitos (?), 1963, L. Moore 50 (holotype US!).

Leme, Elton M. C., Zizka, Georg, Paule, Juraj, Santoro, Julián Aguirre-, Heller, Sascha, Ramírez-Morillo, Ivón M., Halbritter, Heidemarie, Mariath, Jorge E. A., De, Jordano D. T., Forzza, Rafaela C. (2021): Re-evaluation of the Amazonian Hylaeaicum (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular evidence. Phytotaxa 499 (1): 1-60, DOI: 10.11646/phytotaxa.499.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.499.1.1

Hylaeaicum levianum (L.B. Sm.) Leme & Forzza, comb. nov. Basionym:— Neoregelia leviana L.B. Sm., Phytologia 16: 460, pl. 1, fig. 5. 1968. Type:— BRAZIL. Amazonas: Rio Caubouris, 1967, M. Mee s.n. (holotype US!)., Published as part of Leme, Elton M. C., Zizka, Georg, Paule, Juraj, Santoro, Julián Aguirre-, Heller, Sascha, Ramírez-Morillo, Ivón M., Halbritter, Heidemarie, Mariath, Jorge E. A., De, Jordano D. T. & Forzza, Rafaela C., 2021, Re-evaluation of the Amazonian Hylaeaicum (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular evidence, pp. 1-60 in Phytotaxa 499 (1) on page 48, DOI: 10.11646/phytotaxa.499.1.1, http://zenodo.org/record/5424411

Hylaeaicum tarapotoense (Rauh) Leme, Zizka & Aguirre-Santoro, comb. nov. Basionym:— Neoregelia tarapotoensis Rauh, Trop. Subtrop.Pflanzenw. 52: 68. 1985. Type:— PERU. San Martin:via Tarapoto-Yurimaguas, 1200 m elev., 17 August 1983, W. Rauh & Bismarck 63976 (holotype HEID!)., Published as part of Leme, Elton M. C., Zizka, Georg, Paule, Juraj, Santoro, Julián Aguirre-, Heller, Sascha, Ramírez-Morillo, Ivón M., Halbritter, Heidemarie, Mariath, Jorge E. A., De, Jordano D. T. & Forzza, Rafaela C., 2021, Re-evaluation of the Amazonian Hylaeaicum (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular evidence, pp. 1-60 in Phytotaxa 499 (1) on page 49, DOI: 10.11646/phytotaxa.499.1.1, http://zenodo.org/record/5424411

Hylaeaicum eleutheropetalum var. bicolor (L.B. Sm.) Leme & Forzza, comb. nov. Basionym:— Neoregelia eleutheropetala var. bicolor L.B. Sm., Phytologia 8: 227. 1962. Type:— PERU. Loreto: near Loreto, 1961, L. Moore s.n. (holotype US!, isotype NY!)., Published as part of Leme, Elton M. C., Zizka, Georg, Paule, Juraj, Santoro, Julián Aguirre-, Heller, Sascha, Ramírez-Morillo, Ivón M., Halbritter, Heidemarie, Mariath, Jorge E. A., De, Jordano D. T. & Forzza, Rafaela C., 2021, Re-evaluation of the Amazonian Hylaeaicum (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular evidence, pp. 1-60 in Phytotaxa 499 (1) on page 48, DOI: 10.11646/phytotaxa.499.1.1, http://zenodo.org/record/5424411

Hylaeaicum pendulum (L.B. Sm.) Leme, Zizka & Aguirre-Santoro, comb. nov. Basionym:— Neoregelia pendula L.B. Sm., Phytologia 9: 244, pl. 1, figs. 4, 5. 1963. Type:— PERU. Amazonas: Província Bagua, rainforest along Rio Santiago, 3-5 km above mouth, 250-300 m elevation, 8-13 October 1962, J.J. Wurdack 2166 (holotype US!)., Published as part of Leme, Elton M. C., Zizka, Georg, Paule, Juraj, Santoro, Julián Aguirre-, Heller, Sascha, Ramírez-Morillo, Ivón M., Halbritter, Heidemarie, Mariath, Jorge E. A., De, Jordano D. T. & Forzza, Rafaela C., 2021, Re-evaluation of the Amazonian Hylaeaicum (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular evidence, pp. 1-60 in Phytotaxa 499 (1) on page 48, DOI: 10.11646/phytotaxa.499.1.1, http://zenodo.org/record/5424411

Hylaeaicum margaretae (L.B. Sm.) Leme & Forzza, comb. nov. Basionym:— Neoregelia margaretae L.B. Sm., in Mee, Flowers of the Brazilian Forests Pl. 25. 1968. Type:— BRAZIL. Amazonas: Rio Içana, tributary of upper Rio Negro, January 1965, M. Mee s.n. (holotype US!)., Published as part of Leme, Elton M. C., Zizka, Georg, Paule, Juraj, Santoro, Julián Aguirre-, Heller, Sascha, Ramírez-Morillo, Ivón M., Halbritter, Heidemarie, Mariath, Jorge E. A., De, Jordano D. T. & Forzza, Rafaela C., 2021, Re-evaluation of the Amazonian Hylaeaicum (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular evidence, pp. 1-60 in Phytotaxa 499 (1) on page 48, DOI: 10.11646/phytotaxa.499.1.1, http://zenodo.org/record/5424411

Hylaeaicum mooreanum (L.B. Sm.) Leme, Zizka & Aguirre-Santoro, comb. nov. Basionym:— Neoregelia mooreana L.B. Sm., Phytologia. 8: 227, pl. 2, figs. 6-8. 1962. Type:— PERU. Loreto: Vicinity of Iquitos, upper reaches of the Napo River, 1960, L. Moore s.n. (holotype US!)., Published as part of Leme, Elton M. C., Zizka, Georg, Paule, Juraj, Santoro, Julián Aguirre-, Heller, Sascha, Ramírez-Morillo, Ivón M., Halbritter, Heidemarie, Mariath, Jorge E. A., De, Jordano D. T. & Forzza, Rafaela C., 2021, Re-evaluation of the Amazonian Hylaeaicum (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular evidence, pp. 1-60 in Phytotaxa 499 (1) on page 48, DOI: 10.11646/phytotaxa.499.1.1, http://zenodo.org/record/5424411

O uso de caracteres embriológicos tem-se mostrado promissor para relacionar subfamílias, tribos, gêneros e espécies de Rubiaceae. A partir da ginosporogênese (megasporogênese) e formação do rudimento seminal (óvulo) de Borreria verticillata (L.) G. Mey. este trabalho visa enquadrar a espécie nas propostas evolutivas já existentes para Rubiaceae, além de disponibilizar informações para a taxonomia do gênero. A espécie possui ovário ínfero, bilocular e bicarpelar, com um rudimento seminal por lóculo, que, em seu desenvolvimento completo, é anátropo, unitegumentado e tenuinucelado, disposto em placentação axial. O primórdio seminal possui três camadas: dérmica, subdérmica e central. O início da formação do rudimento se dá por divisões não preferenciais da camada central. Em seguida, células derivadas da camada dérmica originam o tegumento e a epiderme nucelar. O arquespório é composto por uma a cinco células arquesporiais e a epiderme nucelar por duas a três células. Do conjunto de células arquesporiais, de uma a três se desenvolvem em células-mãe de ginósporos (células-mãe de megásporos), cuja meiose origina uma tétrade linear, sendo funcional o ginósporo (megásporo) calazal. Comparando os resultados obtidos com a literatura especializada enquadra-se B. verticillata no tipo Mitracarpus, devido à presença de remanescentes do tegumento externo, inclusão do nucelo na calaza, ao número reduzido de células alongadas na epiderme nucelar e ao incremento das células arquesporiais. The structural characteristics of the ovule formation are an important tool for the systematics of the Rubiaceae family. This paper describes the ovule ontogeny and ginosporogenesis (megasporogenesis) of Borreria verticillata (L.) G. Mey. in order to classify this species based on the evolutive diagrams proposed for the Rubiaceae family. The studied species presents an inferior, bicarpellate, bilocular ovary, with one ovule per loculus. The ovule is anatropous, unitegmic, and tenuinucellate, with axial placentation. The ovule primordia present three distinct layers of cells: dermal, subdermal and central. The integument, nucellar epidermis and ovule primordia originate respectively by divisions in the dermal, subdermal and central layers. The archesporium is constituted by one to five cells, while the nucellar epidermis has two to three cells. One to three ginospore mother cells, originated by divisions in archesporial cells, undergo meiotic divisions developing a linear tetrad. Only the chalazal ginospore remains and becomes functional. The remnant cells from the external integument, the inclusion of the nucellus in the chalaza, the occurrence of few elongated cells in the nucellar epidermis and the increased number of archesporial cells classify the ovule formation of B. verticillata in the Mitracarpus type.