Taxonomy of Hohenbergia lanata Pereira & Moutinho, new collections with an amendment to its description

: Hohenbergia lanata Pereira & Moutinho is an endemic bromeliad of the State of Bahia (Brazil), occurring in an altitudinal transition forest between Caatinga dry woodland and Atlantic Forest. The type material was collected by Roberto Burle-Marx, who introduced it into cultivation in his gardens, but lacking the accurate provenance, only mentioning that it was collected between Milagres and Amargosa municipalities. It was later described by Pereira & Moutinho based on a cultivated specimen. In this work, we present new specimens found in natural populations in an area that coincides with the type locality description, enabling us to expand the description of the species, as well as to provide images of the living plants in the field, adding information about its habitat. The herbarium collections revision and the new population found in the field enabled us to propose an updated conservation status and to better characterize the habit, plant architecture, and to add new morphological comparisons and geographical distribution.

Hohenbergia lanata Pereira & Moutinho (1980:88) might be considered among the rarest species of the genus considering the limited collections and the small occurrence area.The addition of morphological data to the description of the species, based only on the holotype, is important, since morphological features, such as the petal description, as well as the morphological variation for measurements of several traits, are limited.Additionally, the species was illustrated only by the image of the holotype, lacking images of living plants.Only a few herbarium specimens of this species were collected after its discovery and it was also reported in the field notes of an expedition to Bahia by Leme (1986).The type specimen was collected between the municipalities of Amargosa and Milagres in Bahia, without an accurate provenance of the type locality or the type of environment in which the species grows.The species can be recognized by its deep tubular rosette, composed of few ligulate and erect leaves covered by a dense white-lanate indument, by the pinkish inflorescence, also covered by a dense white-lanate indument, by its green and strobilate spikes, and flowers with yellow petals, which is a rare feature for Hohenbergia.The use of the specific epithet "lanata" by the authors refers to the dense whitelanate indument that covers both the vegetative body and the inflorescence (peduncle, lateral peduncle, branches, and flowers).This species was initially collected by the Brazilian landscape designer Roberto Burle-Marx, and cultivated in his personal greenhouse, and described by Pereira & Moutinho (1980) when the specimen bloomed.
As part of an ongoing project about the distribution and reproduction of the genus Hohenbergia, in this article we provide an amendment to the description of H. lanata to complement some important traits in the protologue description, as well as morphological variations to all characteristics.We also provide a new morphological comparison with similar species, comments on distribution and habitat, and provide its current conservation status.

MATERIALS AND METHODS
Hohenbergia lanata and similar species (H.correia-araujoi and H. ramageana) were collected during field works at Bahia state (Brazil) in the region of the Caatinga and Atlantic Forest, from 2018 to 2019.The morphological description of the species was based on living plants and in the holotype, and the measurements were taken during field expeditions and from plants maintained in the greenhouse, examined in the Herbarium of Universidade Federal do Recôncavo da Bahia (HURB), Embrapa Mandioca e Fruticultura, and Centro de Energia Nuclear na Agricultura (CENA-USP).Additionally, the species was searched in online databases, namely SpeciesLink (http://splink.cria.org.br/) and Jabot (http://jabot.jbrj.gov.br/v3/consulta.php), resulting in nine specimens from the HURB, ALCB, HUEFS, and RB collections.For the taxonomic description, all specimens deposited in the last 20 years were considered.The conservation assessment for the species was based on geographic distribution and analyzed using the GeoCat software (geocat.kew.org) applying the IUCN red list category criteria (IUCN 2014) and taking into account all the nine specimens verified by the first author and listed as additional examined material.
Type: BRAZIL.Estado da Bahia, entre Milagres and Amargosa, R. Burle Marx s.n., fl.cult.dez.1979.Holotype HB69482!Amended description: Plant epiphytic or rarely terrestrial, 150-250 cm tall when flowering, tubular rosette (small distance between leaf sheaths limiting the capacity to impound water), tank ca.50x30 cm tall, propagates by basal shoots, stem short.Leaves <10 in number, up to 150 cm long, coriaceous, covered by a dense white-lanate indument (the indument follows the leaf venation), erect for ca.2/3 of the height and then slightly recurved, forming a deep tubular rosette impounding water; sheath ca.20 x 14-15 cm, ovoid to elliptic, smooth margins, brownish white to whitish indument on both surfaces, highly coriaceous; leaf blade 80-130 x 4-7 cm, linear to lanceolate, vinaceous or greenish (full sun grown plants can show yellowish leaves) with the adaxial surface lustrous, margins densely serrate, with hard spines (less than 2 mm long), similar to the leaf blade in color, apex acute to acuminate and erect, rarely slight reflexed (apical mucro ca. 1 cm long) or rounded.Peduncle 55-145 cm long, ca.1.5 cm diam., erect, light pink along all its extension and covered by a dense whitelanate indument; peduncle bracts 10-21 x 1.1-1.8cm, lanceolate with an acuminate apex, smooth margins, light brownish, chartaceous, imbricate, exceeding and hiding the internode and covered by a dense white-lanate indument; inflorescence (fertile portion) 60-85 cm long, 28-40 cm diam., commonly 3-branched at the base, laxly pyramidal, erect to slightly suberect, covered by a dense white-lanate indument; primary bracts of basal branches 10-17 x 1.1-1.3cm, lanceolate with an acuminate apex, smooth margins, shorter than branches in length, papyraceous, light brownish; basal branches 12-19 cm long, longstipate (lateral peduncle of basal branches 7-13 cm long) commonly 3-branched, light pinkish covered by a dense white-lanate indument; primary bracts of the middle branches 5-9 cm long, similar to the basal bracts in shape and color; middle branches 8-17 cm long, shortstipate or sub-sessile the lateral peduncle 4-11.4 cm long, similar to the basal branches in shape and color, but 2-branched; primary bracts of distal branches 2-3.5 cm long, similar to the basal bracts in shape and color; distal branches 6-8 cm long, sessile, with 5-8 spikes congested in the apical portion of the rachis, similar to the basal branches in shape and color; spikes 6-8 cm long, 1-1.5 cm diam., cylindrical, greenish and covered by a dense white-lanate indument, presenting up to 50 flowers per spike.Flowers 1.9-2.1 cm long, tubulate, sessile, flattened in the dorsal portion, spirally arranged on the branch, diurnal with a discreet sweet odor; floral bracts 1.4-1.6 x 1.2-1.3cm, triangular to slightly suborbicular, concave with a slightly reflexed apex, apex pungent, margins smooth, greenish, and covered by a dense white-lanate indument, commonly reaching the same height as the sepals; sepals 1.1-1.3x 0.8 cm, triangular with an acute apex, partially free from each other, with lateral wings on both sides that exceed the mucro, margins smooth, whitish; petals 1.6-1.8cm long, distinctly yellow, erect and spatulate, free from each other, with the petal appendage reaching half of the length of the petal; stamens included, 1.3-1.4cm long, shorter than the petals; filaments 1.1-1.2cm long, whitish, partially adnate to petals; anthers 0.2-0.3cm long, yellowish; style ca.1.4 cylindrical, whitish; stigma conduplicate-spiral, higher than anthers and shorter than the petals; ovary ca.0.3 x 0.5 cm diameter, slight pyramidal, dorsal side rounded, axile placentation restricted to the apical portion of the locule; ovules numerous, caudate.Fruit berry, ca. 2 cm long, cylindrical to pyramidal, whitish, indument persistent, blue Bahia, Milages, Serra do Jatobá, Costa, G. 3624, Jul 2018 (UFRN25954).
Distribution and ecological notes: Hohenbergia lanata is an epiphytic or terrestrial bromeliad that grows only in altitudinal transition areas (ecotonal) between Caatinga and Atlantic Forest, preferring high insolation, but also growing near shrubs and trees, showing a disjunct distribution throughout the mountain range of the region of Milagres (12°55'50.1"S39°47'21.9"W)(Fig. 3).The main population found for H. lanata is in the upper region of Serra do Jatobá, placed along the highway that connects Amargosa to Milagres, 12º55'24"S; 39º46'44"W, at an elevation of 764 m.s.m., near the base of the mountain range.There, it is common to observe the species growing epiphytically on palm tree stems (Licuri, Syagrus coronata), creating dense clumps (Fig. 1).Small and fragmented populations of H. lanata are also observed in other mountain ranges in the region, such as Santa Teresinha municipality (12°50'53.5"S39°28'29.7"W,starting at 750 m.s.m), Serra da Jiboia (localized at highway BA120, between Santa Teresinha and Elísio Medrado municipalities,) and in Fazenda Ferrugem (Elísio Medrado municipality, at highway BA026, which connects Amargosa to Santo Antonio de Jesus, 12°58'42.4"S39°31'05.0"W).The description of deposited specimens indicate that the species can also grow in other areas within the Amargosa city regions, such as Ibipitanga, but always in altitudinal caatinga.The leaves are predominantly vinaceous, with a few plants presenting greenish leaves.
Phenology: The blooming season is between January and April, during the rainy season.The herborized collections indicate that some plants may also bloom between July and September.
Conservation status: The main locality of the species is surrounded by a busy federal highway and the other two localities are natural areas of extractivism, making the species vulnerable to extinction.By plotting the known occurrences in GeoCat software (geocat.kew.org), it was estimated that the extent of occurrence is about 1,094.202km 2 while the estimated area of occupancy is 20 km 2, defining the species as Endangered (EN).Since these areas are under threat of a rapidly human expansion and crossed by high traffic roads, we categorized the species as Endangered (EN), following the IUCN definition (IUCN 2014).We reinforce that it is only known from small fragments of forests, where the quality of the habitats can decrease rapidly, despite one of these localities being a private reserve.
Remarks: Hohenbergia lanata was compared with H. ramageana Mez (1896:127) by Pereira & Moutinho (1980), mainly due to questionable characteristics presented by Smith & Down (1979), such as plant and inflorescence shape, leading the authors to compare two plants with completely different shapes and habits (Baracho 2004).Baracho (2004) also related H. lanata with H. ramageana and explained that both differed only by the size of sepals and spikes.Currently H. ramageana has a better circumscription compared to other Hohenbergia species from the Atlantic Forest, it presents a crateriform rosette formed by patent leaves (while species from Caatinga habitat have a tubular rosette and erect leaves, as also observed in H. lanata) and pyramidal and 4-branched long-stipate inflorescence with short spikes (vs.pyramidal, single-branched short stipate inflorescence with long spikes in H. lanata).Additionally, these species show many morphological differences, as shown in Table I.
In this work, we compared H. lanata with H. correia-araujoi Pereira & Moutinho (1980:88), both endemic to Caatinga forests of Bahia, sharing many characteristics, including the shape of the plant and the ecological characteristics (Table I).Initially, it is important to emphasize that H. lanata and H. correiaaraujoi present unique plant and inflorescence architectures within the genus.The presence of a tubular rosette with long leaves (> 80 cm long) and a triangular inflorescence distinguish both of them from Campos Rupestres species (which have lageniform rosette with short leaf blades and a sub-cylindrical inflorescence).Additionally, the tubular rosette, the erect leaves, and the combination of the light pink inflorescence covered by a dense white-lanate indument, light green floral bracts, and yellow petals distinguish them from the Atlantic Forest species (which have patent leaves forming a crateriform rosette, reddish or green inflorescence covered by a brownish or white indument, red or yellowish floral bracts and purple or white petals) (Smith & Down 1979, Baracho 2004).
Comparing Hohenbergia lanata with H. correia-araujoi, H. lanata is identified by the absence of the horizontal scale bands in the leaves (unique to H. correia-araujoi) however, additional discrete traits can also be indicated.Hohenbergia lanata presents a constrict rosette (the spaces between the sheaths are smaller than in H. correia-araujoi, creating a deep tubular rosette), longer and narrower leaves (up to 150 x 4 -7 cm vs. less than 70 x 12 -15 cm in H. correia-araujoi), acuminate leaf apex (vs.cuspidate to acute leaf apex), dense white-indument on both leaf surfaces with no ornamentation (vs.discreetly

Figure 1 .
Figure 1.Hohenbergia lanata.a) An epiphytic population in full bloom.b) A blooming plant.c) Details of the upper portion of the inflorescence, showing branches and one flower in anthesis (arrow).d) A terrestrial plant and inflorescence in detail.

Table I .
Morphological comparisons between Hohenbergia lanata, Hohenbergia correia-araujoi, H. ramageana,  and H. flava.The measurements are based on the original description of each species and were complemented by observations of living plants.indumentwithhorizontalscalebands ornamentation), longer basal branches (up to 19 cm long vs. less than 14 cm long), bracts of the lateral peduncle shorter than branches (vs.longer than branches).Another Hohenbergia species that lives near Hohenbergia lanata is H. flavaLeme (2004:22), both species present a quite lanate inflorescence with greenish branches and yellow petals, but H. flava can easily be distinguished by its crateriform rosette with patent and yellowish leaves, inflorescence with a sessile branch, and sharp floral bract.Other characteristics of the inflorescence are very similar when both are compared, thus a detailed morphological comparison among H. lanata, H. correia-araujoi, H. ramageana, and H. flava is presented here (TableI). brownish