Vellozia hirsuta and V . tubiflora ( Velloziaceae )

(Circumscribing Vellozia hirsuta and V. tubiflora (Velloziaceae)). Vellozia hirsuta and V. tubiflora belonged to Vellozia sect. Radia, a poliphyletic taxon. Due to different choices of characters for delimiting the species, from 21 species formelly included in the section, only three are now recognised. The hypanthial tube longer than ovary, and deciduous in fruit, distinguishes V. hirsuta and V. tubiflora from all other Velloziaceae. Although consistently delimited, they exhibit wide intraspecific variation. Thus, V. hirsuta encompasses four taxonomic synonyms, and V. tubiflora, 19. On the other hand, the many different populations of both species lead to the recognition of 14 morphological patterns within V. hirsuta and eight within V. tubiflora, although those patterns cannot receive taxonomic status.


Introduction
Vellozia hirsuta and V. tubiflora belonged to the former Vellozia sect.Radia, defined as being composed of species having flat leaves, with almost always distinct grooves on the abaxial side, tepals fused into a long tube, and more than six stamens (Smith & Ayensu 1976).Of these attributes, the long hypanthium tube is the most characteristic and has also been used by Menezes (1980) to define the section.Cladistic analyses (Behnke et al. 2000, Mello-Silva 2000, 2005) suggested that Vellozia sect.Radia should be a polyphyletic group.In Mello-Silva (2000, 2005) V. hirsuta and V. tubiflora emerged as closed related species, although in Mello-Silva et al. (unpublished data) they belong to rather distinct clades.From a total of 21 species admitted to that section by Smith & Ayensu (1976), only four were recognized by Mello-Silva (2000).These contrasting positions reflect different opinions about the aspects to be addressed in the delimitation of the species in Velloziaceae.Non-variable characters have often been ignored in favour of others more prone to variation.Smith & Ayensu (1976) used the indumentum, the dimensions of the pedicel, and the hypanthium tube as the basis to distinguish the various species within the section Radia, although these characters may not be very consistent with species limits, nor with different populations of the same species (Mello-Silva 1990, 1995).An example of this situation is the synonymization of V. leucanthos with V. cachimbensis, by Smith & Ayensu (1976), based on similarities such as habit and vestiture, while neglecting characters such as aquiferous parenchyma and phloem strands, which are now used to merge those species into V. hirsuta and V. tubiflora, respectively.Their variation is here characterized and discussed in a similar, although much more encompassing, study that has been done for V. hirsuta solely (Mello-Silva 1990).This paper also provides an identification key, descriptions, synonymy, typifications, illustrations, phenology, and geographic distribution of the species.

Material and methods
Morphological studies where conducted using herbarium specimens as well as some spirit collections.Studied material and the herbaria where they are deposited are arranged in geographical order, from North to South.Herbarium specimens not examined are indicated by n.v.(non vidi).Herbarium acronyms follow Holmgren & Holmgren (1998).
Transversal sections of the median portion of the leaf blades and pedicels from dried specimens were made employing a hand-held razor blade.Sections were stained with oyster-blue and basic fucsin according to Roeser (1972) and adapted by Luque et al. (1996).Median portion of leaf blades and seeds were mounted on aluminium stubs, coated with gold, and examined with a scanning electron microscope Zeiss DSM 940.Drawings were done in pen-and-ink, with the aid of a Wild M8 stereomicroscope equipped with a camera lucida.Type specimens (holotypes and/or isotypes) of all names were examined, except of V. markgrafii and V. macrosiphonia, which were not located, and of V. dumitiana.From this last one, paratypes were examined.

Results and Discussion
Characters circumscribing Vellozia hirsuta and V. tubifora are those also used in cladistic analyses (Mello-Silva 2000, 2005).Vellozia hirsuta is delimited by one phloem strand, and solitary flowers; V. tubiflora by smooth subsidiary cells in stomata, aquiferous hypodermis extending to bundle sheaths and furrows, spiral thickenings in exotesta of seeds absent (Sousa 2005), and a sclerified belt in pedicels.The hypanthial tube always longer than ovary, and deciduous in fruit, distinguishes both species from all other Velloziaceae.Although consistently delimited, V. hirsuta and V. tubiflora exhibit a wide infraspecific variation.They comprehend many different and more or less isolated populations, distributed over a large area, and genetic exchange between populations is very improbable.The populations are found on isolated mountains, what may have allowed the appearance and fixation of distinctive characteristics in each group, increasing their diversity.These differences occur in a reticulated pattern within the populations, and leads to the recognition of two complex species.The only other option would be their fragmentation into a number of species, a majority with a restricted distribution, whose delimitations would be problematic, if not impossible.Plants caespitose or solitary.Stem 1.5-120 cm long, 0.4-5 cm diam.at apex, entire or few-branched, rarely much-branched.Living leaves 3-16 in each branch, tristichous.Leaf sheaths adaxially glabrous, abaxially densely lanate to lanate-tomentose to glabrescens, or glabrous at the base and glabrescens at the apex; apex curved or straight, entire or lacerate.Leaf lamina (2-)6-37 cm long, 1.5-15 mm broad, linear-triangular, plane or arcuate, twisted, in sicco either revolute or conduplicate near midrib and plane near margins, at least at distal portion, rarely involute, erect or erect to spreading, green, deciduous or marcescent and reflexed, densely to sparsely covered with trichomes of varying lengths up to 1 cm, these flat, entire, branching or in fascicles, larger and more densely disposed on the abaxial surface and towards the base, otherwise the lamina lanate on the margins and on central nerve at abaxial surface or glabrescens or glabrous; either very viscous to viscous to not viscous at all, odoriferous or not; margins entire, hardly to clearly thickened, indumentum similar to that of the lamina; apex generally long, rarely shortattenuate.Flower 1 per branch, odoriferous.Pedicel 1-21 cm long, 1-3 mm diam., trigonous, shorter than the leaves, rarely longer (Mello-Silva 454), evident, sometimes hidden amongst the leaves and sheaths, and then up to 4 cm long, the flower appearing sessile; during fructification either of the same length or expanded up to 7.5 cm long and c. 4 mm diam., green, lighter and smooth towards base, bearing small glandular emergences in the apical half, sometimes totally smooth.Hypanthium, in the ovary region, oblong-trigonous, with attenuate angles, 5-20 mm long, 2-7 mm broad, green, densely covered with emergences, these glandular-stipitate with concave, truncate, umbonate or subulate apex, sometimes subulate or densely tomentose.Hypanthial tube 2.5-11 cm long, 1-4 mm diam.at base, 2-7 mm diam.at apex, white to slightly purplish, green or greenish purple at base, with emergences similar to those of the ovary region but smaller and sparsely distributed at the base and the region continuous to the external tepals.Tepals 4.5-7.5 cm long, 1-3 cm broad, oblong-elliptical, arched and spreading in the apical half, white, rarely purple, the outermost bearing emergences similar to that of the hypanthial tube on the proximal abaxial surface and on the midrib, adaxial surface glabrous; the innermost glabrous on both surfaces or with the emergences at the base of abaxial surface.Stamens 9-24, all equal, exserted from tube; filaments c. 2.5-3 cm long, white to greenish; anthers 1-3 cm long, yellow to pale-yellow, latrorse; staminal appendages absent.Style 5-12.5 cm long, white to purplish-white, protruding beyond the stamens; stigma 3-6 mm diam., yellow to white.Capsule 1-3.3 cm long, 6-25 mm diam., ellipsoid to oblong-trigonous to suborbicular-trigonous to spheroid, loculicidal, green to greenish-purple when immature, chestnut brown at maturity, at first covered by the hypanthium, with emergences like those of the flower, thicker or not, the hypanthium later caducous.Seeds numerous, 2-3 mm long, cinereous, exotesta with spiral thickenings.Foliar and pedicellar anatomy (figures 1A, C, G, I, 3) -Lamina dorsiventral.Furrows c. 1/3-1/4 of the thickness of the lamina, rarely absent, smooth.Cuticle thicker on the adaxial surface.Adaxial epidermis pluriseriate (types 1, 2 and 3) or uniseriate (types 4 and 5), abaxial uniseriate.Stomata always present on the abaxial surface, usually inside the furrows; on the adaxial surface present either on entire surface (type 5), or only on the inner half of lamina (types 2, 3 and 4) or only near the midrib to absent (type 1).Aquiferous hypodermis of the adaxial surface 1-2-seriate, extending to endodermis as aquiferous parenchyma.Palisade parenchyma 3-4 cell layers thick, abaxially merging with spongy parenchyma.Fibro-vascular bundles surrounded by endodermis as leaf bundle sheath.1-2 larger vessels present in each bundle.Phloem strand 1, V-shape beneath the xylem.Pericyclic fibres extending adaxially to the aquiferous parenchyma and abaxially to (or nearly to) the hypodermis (in Mello-Silva 508, laterally enlarged at base, reaching the furrows [figure 1C]).Fibres strands (1-)3-6(-10) cell layer thick on all adaxial surface (type 1) or (1-)2-3 cell layer thick Dentro, Menezes 344 (BHCB, SPF); Without precise location: cultivated on Rio de Janeiro, Menezes 780 (SPF), 903 (SPF), 1225 (SPF).
Vellozia hirsuta occurs in the Espinhaço range in Brazil, from the northern part of the Serra do Cipó to Espinosa, in Minas Gerais State.Some disjunct populations occur near Catas Altas, Jequitinhonha and Pedra Azul, in Minas Gerais, and near Cordeiros, Caetité, Macaúbas, and Jequié, in Bahia State.This species is found either in the high mountains, covering large rocky surfaces, or on smaller rock outcrops in the midst of arboreal or shrub vegetation in regions of cerrado and caatinga vegetation (figure 2).
Mello-Silva (1990) recognized the existence of six patterns in Vellozia hirsuta, in a combination of five diferent types of leaf anatomy (figure 3) and three types of vegetative morphology.Nevertheless, those anatomical types are not fixed.Analyses of 4,500 leaves from approximately 800 individuals from 18 populations (table 1) reveal intermediate types between anatomical types 1 and 2 (figure 3C) and between anatomical types 3 and 5 (figure 3K).These variations occur principally in Grão-Mogol, where populations with different types are in close contact.Additionally, two nearby populations in Diamantina demonstrate a certain degree of variation.
Populations showing type 1 appear to be less subject to variation, while type 5 populations are the most variable.This may be due to the greater isolation of the majority of type 1 populations.In addition, one population with plants of type 5 (Mello-Silva 454), which is geographically isolated, demonstrates a low level of variation, suggesting that greater variation is the result of greater genetic flux among populations.Within the same leaf, the type of leaf anatomy is always uniform (figure 3A, B, H-J) although different leaves on the same individual can show different types.Two plants from Mello-Silva 465, which showed the highest degree of variation, had all their leaves examined.The first of these plants had four leaves with anatomical type 3, nine leaves that had types intermediate between 3 and 5, and seven leaves having type 5.The second individual had three leaves with intermediate types between 3 and 5, and seven leaves with type 5. Another leaf from Windisch 337, the only collection which was found to possess anatomical type 4 (Mello-Silva 1990), showed one fibre bundle in the adaxial epidermis, characterizing a intermediate type between 2 and 4.
Thus, there are populations that have anatomical types and external morphology intermediate between the major patterns established by Mello-Silva (1990), and whose geographical distribution indicates a reticulated pattern of interrelations.Fourteen patterns of variation can be recognized, which are reasonably distinct, although non-discrete.These patterns could be distributed into two more or less distinct and sympatric assemblages.One of these assemblages (patterns 1 to 5, bellow), which has been considered as a population complex of V. hirsuta (Mello-Silva 1990, 1995) In the first assemblage, which includes the types of V. hirsuta, V. leucanthos, V. maguirei, and V. riedeliana, the dried leaves are revolute, rarely involute; the leaf blade is viscid and generally odoriferous, almost always without a well defined abscission zone; the flowers are more delicate, almost always with an evident pedicel; the hypanthium emergences are short and truncated, rarely subulate, and never covered by a tomentose indumentum.In contrast, the plants from the second assemblage, which includes the type of V. markgraffii, have leaves conduplicate near their centre and flat along the margins, rarely revolute (Mello-Silva 629), and generally more sclerified (figure 1C); neither viscous, nor odoriferous, with abscission zone almost always well defined; flowers more robust, with their pedicel almost always hidden; and hypanthium emergences either cylindrical with truncated apices, concave or umbonate, or long-subulate, often covered with a tomentose indumentum.These assemblages, whose complexity will be described below, could be classified into two distinct species (Mello-Silva 1995) were it not for the lack of universal characteristics that would allow one to distinguish them, as well as for the existence of a number of intermediate populations  285,391,466,473,484,515,526,1421,1453,1501,2559,2698,3176,3196,3202,3222       Solitary, rarely caespitose, plants.Stem 2-200 cm long, 0.5-5 cm diam.at apex, entire to much-branched.Living leaves 4-17 in each branch, tristichous.Leaf sheaths adaxially glabrous, abaxially densely lanate or floccose or villose-tomentose to glabrescens or glabrous; apex recurved or straight, entire or lacerate.Leaf-lamina 6.5-65 cm long, 3-22 mm broad, linear-triangular, arcuate or plane, in sicco twisted and revolute, often straight or involute, green to cinereous-green, sometimes purplish when young, erect to erect-patent, deciduous, rarely some marcescent and reflexed, abaxial side densely to sparsely sericeo-to hirsute-lanate, trichomes of varying lengths, stellate-rotate, dendritic-spatulate or dendritic-filiforms, erect to adpressed, adaxial side mostly glabrous, sometimes with indumentum similar to that of the abaxial side, but less developed and sparsely distributed, rarely of the same appearance; lamina sometimes glabrous on both sides or ciliate on margins and midrib on abaxial side or ciliate only in the margins or lamina pubescent on both sides and glabrescens on the adaxial side; rarely viscous and/ or odoriferous; margins with indumentum similar to that of the lamina or ciliate; apex attenuate.Flowers 1-7(-15) per branch, odoriferous.Pedicel 4.5-24 cm long, 1-2 mm diam., trigonous, lightgreen, somewhat purplish towards the base and apex, sometimes completely purple to dark-purple, evident, smooth towards the base, bearing glandular emergences towards the apex.Hypanthium, in the ovary region, oblong-trigonous with attenuate angles, 5-15 mm long, 2-5 mm diam., green to greenish-purple, sometimes dark-purple, densely covered with glandular emergences, truncate or not, with 1-4 mm long.Hypanthial tube 2.3-10.5 cm long, 1-4 mm diam.at base and 2-5 mm diam.at apex, mostly white, sometimes green or greenish-purple or reddish or dark-purple at base, or totally purple, with glandular emergences sparsely distributed at the base and the region continuous to the external tepals, deciduous at fructification.Tepals 2.5-7.5 cm long, c. 1-2 cm broad, oblong-elliptical, mainly white, sometimes violet at base and midrib or totally violet to deep purple, outermost bearing small glandular emergences at base and midrib at abaxial side, adaxial side smooth; the innermost glabrous.Stamens 12-18, all equal, exserted from tube; filaments c. 2-15 mm long, white; anthers c. 1-2 cm long, yellow, latrorse; staminal appendages absent.Style 5-12.5 cm long, white, protruding beyond the stamens; stigma 2-7 mm diam., yellow.Capsule 1-2.3 cm long, 0.8-1.8cm diam., oblong-ellipsoid to oboval-oblong, trigonous, loculicidal, green or greenish-purple when immature, chestnut brown at maturity, at first covered by the hypanthium, this later caducous.Seeds numerous, c. 1.5-2 mm long, verrucose or foveate, dark-brown to blackish, exotesta without spiral thickenings.Foliar and pedicellar anatomy (figures 1B, D-F, H, J, 10) -Lamina dorsiventral.Furrows ca.1/2 of the thickness of the lamina, rarely less deep to inconspicuous, smooth or with more or less conical, small papillae, not very conspicuous.Cuticle thicker on the adaxial surface.Adaxial epidermis (uni-) pluriseriate, abaxial 1-2-seriate.Stomata mainly inside the furrows or on their region in abaxial surface when the furrows are inconspicuous, few on the adaxial surface.Aquiferous hypodermis on the adaxial surface 1(-2)-seriate, extending adaxially, as aquiferous parenchyma, to endodermis and to furrows or to the spongy parenchyma on the region of the furrows when these are inconspicuous.Palisade parenchyma 2-5-cell layer thick, rarely absent (Idobro 632), abaxially merging with spongy parenchyma.Fibro-vascular bundles surrounded by endodermis as leaf bundle sheath.1-2(-3) larger vessels present in each bundle.Phloem strands 2, U-shaped beneath the xylem, protophloems separated by fibres or parenchymatic cells.Pericyclic fibres extending adaxially to the aquiferous parenchyma and abaxially to (or almost to) the epidermis.Fibres strands 1-6 cell layer thick present on adaxial surface or absent and, on abaxial surface, 1(-2-3) cell layer thick present on all surface or near furrows edges or, rarely, absent.Pedicel triangular in transverse section.Fibrovascular bundles 6. Belt of sclerified cells present.Specimens examined: PANAMA.Chiriquí: cerro Vaca, Pittier 5352 (F, US); VeraGuas: La Yeguada, cerro Altos de La Gallota, Correa A. 1911 (MO, PMA n.v.) North of the equator, the area of distribution of V. tubiflora extends over the entire region of the Roraima formation, including the sedimentary mountains of Guyana and the "gran sabana" of Venezuela, the peaks along the frontier with Brazil and, to the west, the Negro and Guainía rivers, in Colombia, in a series of terraces reaching up to the Macarena Range along the flanks of the Andes.It also occurs in the Orinoco, Casiquiare, Negro and Guainía rivers basins, areas that were most likely contiguous before the up lifting of the high plains (Maguire 1970).The presence of this plant along river margins results in considerable differences in altitude among different Venezuelan populations, which are otherwise located very close to one another.Approximately 150 km separates two populations along the margin of the Orinoco river; one grows at 60 m of altitude, and the other population at 1800 m, at cerro Guanay.Less than 30 km separates a population along the Cunucunuma river, at 180 m, from another at cerro Duida, at 2,400 m above sea level.In Panama, V. tubiflora occurs on the eastern flanks of the cerro Chiriqui, where the vegetation is similar to that seen in the high mountains in Colombia (Simpson & Neff 1985).Isolated populations growing on rock outcrops also exist in numerous spots within the Amazon basin.Within the Espinhaço range in Brazil, where the large majority of species of Velloziaceae are found, V. tubiflora only occurs in Bahia State, in the mountains to the northeast of the Paramirim river and to the north of the São Paulo river, an affluent of the Brumado river (figure 2).The isolated populations of V. tubiflora exhibit morphological and anatomical divergences, principally in terms of their habit and V. duidae.Pattern 2 - (Dawson 14674;Glaziou 22218a;Hatschbach 36322, 53929, 60303;Irwin 12454, 12697, 32227;King 8890;Maguire 32307;Martinelli 3789;Menezes 598;Pirani 1683;Rivadavia Lopes 181;Rosa 1994;Toledo 283; Tillet 43855; figure 7B,  C).Includes the type material of V. machrisiana and V. maculata.They have an erect stem from 7 to 10 mm in diameter, rarely 20 mm, and the leaf sheaths at the apex of the stem are revolute.1368).In some collections (Cavalcanti 1368, Mello-Silva CFCR7386), the dry leaves will remain flat or involute, in contrast to the general state of the species, which show the dry leaves normally as revolute.The leaf sheaths are also woolly and imbricate.The stem is generally between 2 and 3 cm thick, varying in length from 8 to 120 cm.Populations of this pattern are found in Brazil in Serra Dourada, Goiás, in the

Collection Flower colour
Height (cm) Width (mm) Pirineus (pattern 7).Cavalcanti 1342, which has a thin stem and leaves that become revolute when dry, is more similar to pattern 2, except for the indumentum, which is typical of pattern 8.The plants of this pattern are considerably variable in size, which had leaded the authors to recognize two species, V. annulata and V. dawsonii.The first of these had been traditionally defined (Henrard 1937, Smith 1962, Smith & Ayensu 1976) as a species approximately 50 cm tall, with slender stems and leaves, while the second species was more robust, up to 2 meters tall, with much more developed stems and leaves.However, there exists a gradient of variation in relation to this characteristic, with specimens ranging from 10 cm to 2 meters tall, without the existence of discrete categories within this range (Table 2).In relation to other attributes, the collections are identical.In terms of leaf anatomy, the collections exhibit gradation in the quantity and volume of the fibre bundles on the adaxial surface.380, 405, 1327, 1342;Dawson 14580;Duarte 10794;Glaziou 22212, 22217;Harley 11417, 11464;Hatschbach 36314, 36801, 36869, 36908, 60296, 60298;Irwin 12622, 12648, 24344;Mello-Silva 554;Menezes 273, 274, 599, 600, SPF 71864;Oliveira 500;Pereira 1518;Pirani 1824Pirani , 1734Pirani , 1871;;Prance 58275, 58276;Rizzo 8128;Romaniuc Neto 635;figure 9A-H).Includes the types of V. annulata and V. dawsonii.The plants are characterized by flat leaves, which become involute when dried.This is the opposite of the majority of the individuals of other patterns, which have arched leaves that become revolute, or more rarely flat or involute, when dried.
The indumentum is dense and short on the abaxial side of the leaf.Robust or very young individuals may have their basal leaves arched, although the rest of them will be flat.These populations are restricted to Chapada dos Veadeiros in the municipalities of Cavalcante, Alto Paraíso de Goiás and São João da Aliança, Goiás. Cavalcanti 1327, Harley 11464, Hatschbach 36314, 36908, Irwin 24344, Menezes 274 and Pirani 1824 have flat or involute leaves, typical of this pattern, with indumentum similar to that found in pattern 5 and also in the plants from Serra dos Phenology -The large number of collections of V. hirsuta and V. tubiflora furnishes data concerning the modes of flowering and fruiting in these species (table 3).In V. tubiflora, there are two distinct groups with respect to flowering modes.The populations to the north of the equator show a flowering curve practically unaltered during the entire year, while the populations to the south of the equator have a distinct seasonal cycle, with flowering peaks between the months of December and February (table 3, figure 11B).V. hirsuta shows a flowering peak between October and January (table 3, figure 11A).Abiotic factors such as temperature, humidity and photoperiod may be acting in an isolated manner, or together, to produce these phenological patterns.Populations of V. tubiflora to the south of the equator, together with the populations of V. hirsuta, are situated in the South America zonobiome II, with evident seasonality of temperature and humidity, with heavy peak rains in the hot season, and an arid cold season (Walter 1984).These seasonal alterations in the climate seem to generate the type of flowering curves above mentioned, where the flowering peaks correspond to the wettest and warmest months.In contrast, the populations of V. tubiflora to the north of the equator are located for the most part in zonobiome I, in Venezuela and Colombia, or in the transition zone between zonobiomes I and II, in Venezuela, Guyana and Panama, where there is almost no seasonal climatic change (Walter 1984).The uniform climate could well be responsible for the uniformity in the flowering curve.The abiotic factors could influence flowering either directly, by altering the capacity of the plants to produce flowers, or indirectly, by effecting the pollination vectors (Rathcke & Lacey 1985).In Central America, seasonal abundance of butterflies coincids with increased flowering of sphingophilous species.On the other hand, the seasonal increase in butterflies also coincids with production of new leaves, which are necessary for the development of their larvae (Janzen 1967, Frankie et al. 1974, Rathcke & Lacey 1985), suggesting that the pollinator's life cycle is equally influenced by changes in the seasons.As both V. hirsuta and V. tubiflora are probably sphingophilous (Mello-Silva 1995, Sazima 1978, Vogel 1969a, b), this could be a factor effecting their flowering cycles.The influence of the photoperiod must also be considered.
Plants may develop sensitivity to the small annual variations in day length (Daubenmire 1974).At 10° of latitude the days can reach a length of 12 hours and 45 minutes and, at 20°, very near the southern limit of distribution of the species considered here (figure 2), daylight can last for up to 13 hours and 30 minutes (Salisbury & Ross 1992).The species and populations demonstrating seasonal variations in flowering are exactly those found at greater latitudes; the populations of V. tubiflora showing persistent flowering throughout the year are located very near to the equator, where they experience only a small, or essentially nonexistent, differences in photoperiod.Phenological studies of populations situated at the extreme limits of distribution south of the equator (Serra das Flores, Ceará, and Serrana, São Paulo, both in Brazil) could help to clarify the influence of the photoperiod on flowering in these species.
Fruiting immediately follows flowering, thereby generating curves with similar patterns, although with a slight forward temporal dislocation (table 3, figure 11A, C).These results may be a bit imprecise, as the fruits remain attached to the plants for a long period, and their age was not taken into account in this study.Biogeography (figure 2) -The diaspores of V. tubiflora, as well as the majority of the Vellozia, are not adapted for dispersion over long distances.This leads one to believe that V. tubiflora must have had, in the past, an essentially continuous distribution over the continent.Before the Miocene, the flora of the Brazilian and the Guyanan shields was continuous (Maguire 1970).This continuity was interrupted by three consecutive geological events which occurred about 3 million years ago: the rising up of the Andes Mountains, the elevation of the Guyanan plateau, and the resultant change in direction of the flow of the Amazon river (Brown & Gibson 1983).The Amazon at first formed a lake, and then a river system, both of which allowed the installation of an Amazon flora, which served as a barrier between the flora of the mountains of Brazil and that of Guyana (Maguire 1970).Thus, the populations of V. tubiflora in South America were separated from those of Panama, as well as the populations north and south of the line of the equator.Later, the erosion of the highlands and cyclical climatic changes probably served to further fragment these once contiguous populations into three main blocks, as well as many smaller isolated populations.In the Espinhaço range, the fragmentation of an ancient plain by erosion (Abreu 1984) resulted in the isolation of resident populations of V. hirsuta.The absence of V. tubiflora in the greater part of the Espinhaço range is of importance to the biogeography of the region.Its probable ecological correspondent in the region is V. hirsuta.The two species are very similar with respect to their external morphological characteristics, and both are probably sphingophilous (Sazima 1978, Vogel 1969a, b).These similar characteristics have placed them closely related in morphological analyses (Mello-Silva 2000, 2005), but appeared as convergences in a combined analysis (Mello-Silva et al. unpublished data).The area of distribution of V. hirsuta in the Espinhaço range reaches the mountains in Macaúbas, Bahia State.Beyond the Paramirim and São Paulo river valleys, V. hirsuta is no longer found, only populations of V. tubiflora.In no other place they do come closer.In this way, the southern and western portions of the Espinhaço range in Bahia may be more closely related biogeographically to the elevations in Minas Gerais than with the bulk of the Chapada Diamantina mountains that lie further to the northeast.If this statement is correct, the Paramirim, Rio de Contas, and São Paulo river valleys act as vicariant barriers.
Nomenclatural notes -Bellow are present nomenclatural and typification notes of some names involved in taxonomy of both Vellozia hirsuta and V. tubiflora.
Vellozia annulata Goethart & Henrard Goethart & Henrard in Henrard (1937) designated Glaziou 22212 and 22217, mounted on a single sheet in the Leiden herbarium (L), as type collections (syntypi) of V. annulata.Smith & Ayensu (1976) assigned only Glaziou 22212, which is also deposited in BR, K, and P herbaria, as the type collection.In this manner they effected the lectotypification, although they did not convey that.

Vellozia riedeliana Goethart & Henrard
Goethart & Henrard in Henrard (1937) designated Riedel 1051 at LE as holotype of V. riedeliana.Nevertheless, this number is not found at LE and a lectotype should be designated with the isotype housed at L. Vellozia tubiflora (A.Rich.)Kunth Richard used the name Campderia, in the Bull.Soc.Philom. in May of 1822, to describe two New World species of Velloziaceae: Campderia langsdorffii ('Langsderffii') and C. tubiflora.The work is an excerpt, in the third person, of a lecture given by the author to the Société Philomatique de Paris in March of 1822.The name Campderia had already been given to a genus of Umbelliferae (Campderia Lag., Amen.Nat.Españas 2: 99.1821 = Kundemannia Scop.), becoming Campderia A.Rich. a later homonym, and therefore illegitimate (article 53.1, ICBN).Perhaps having become aware that the name had already been used in the Umbelliferae, Richard published, in the Syn.pl. of Kunth in December 9, 1822, Campderia as a generic synonym of Radia (Radia had also been used, in the Sapotaceae, but in this case it was a nomen nudum).If the name Radia was intended as homage to the botanist Raddi, it should be written as Raddia and thus be treated as an illegitimate later homonym of Raddia Bertol.(Bologn. Opusc. Sc. 3: 410. 1891).However, there is no evidence that Radia refers to Raddi and, as such, this generic name can be treated as legitimate.In conclusion, Campderia tubiflora is not the correct name (article 55.1, ICBN) and Radia tubiflora and Vellozia tubiflora should be based on it.In describing Campderia tubiflora, Richard (1822) used material collected by Humboldt and Bonpland on the banks of the Orinoco river, in May 1800.In proposing the combination for Radia tubiflora, Richard (in Kunth 1822) provided a short analysis of the species as well as a detailed description, this being based on a manuscript written by Bonpland.Kunth (1825) used the same procedure when transferring it to the genus Vellozia, noting that the type material had been destroyed (specimina suppetentia carie destructa).This material could not be located in the Paris herbarium (P).Humboldt (1980: 142) noted that many of Bonpland's collections that have been made during May 1800, along the banks of the Orinoco river, were lost due to excess humidity.Thus for the neo-typification of the species, a collection from that same region (Maguire 29245) was chosen.All populations along that river belong to pattern 1 described above.

Vellozia velutinosa Goethart & Henrard
Goethart & Henrard in Henrard (1937) designated Glaziou 22218 as type collection of V. velutinosa, but did not cite any herbaria.Smith & Ayensu (1976) assigned a material in L as the holotype.However, Glaziou 22218 could not be found in that or in any other herbaria, except K, and this has been chosen as lectotype.

Table 1 .
Foliar anatomical types of 4500 examined leaves from 18 populations and of Vellozia hirsuta.
Figure 2. Geographical distribution of V. hirsuta (s) and V. tubiflora (l) in Central and South Americas, showing occurrence of the different patterns (numbers) of each species.Base map modified from Flora Neotropica base map, The State University of Utrecht, the Netherlands.
with golden-hirsute indumentum of very long trichomes (figure4A).These are restricted to Gouveia and Diamantina region, mainly in Biribiri, as the type-collection of the species, Glaziou 19924.The individuals from Rio Vermelho have a very reduced stem and very short trichomes (figure4B).The anatomical type is generally 1 but Giulietti CFCR2376 and Hensold CFCR3184 have anatomical type 2. This pattern has been previously defined as A1 with marcescent leaves (Mello-Silva 1990).
; COLOMBIA.Guainía: vicinity of cerro Monachi, Schultes 10067 (GH); ViChada: Maipures, Orinoco river, Chaffanjon 326 (P); Spruce 3606 (K, P); Thomas 2556 (NY); Raudal San Borja-Ventanas, Pinto E. 1264 (COL, P); 1 km de Casuarito, Pinto E. 1396 (COL n.v., P); El Tuparro, Daniel 138 (COL); The plants are almost glabrous; the trichomes are very short, occurring only on the abaxial side of the sheaths as well as on the margins and on the abaxial surface of the central vein of the leaves.Occasionally, the leaves are glabrous.The leaf blades are slightly arched, almost flat, very similar to, or equal to, the transversal posture of the leaf blade found among members of pattern 8.These plants almost always grow along the banks of rivers or of lesser creeks in the region around Chapada dos Veadeiros, Goiás (e.g.Prance 29160), from the Demini river in Amazonas State, Brazil (Miranda s.n.), and from cerro Guanay, Venezuela (Maguire 31679) have a short and dense indumentum, although in some collections from northern Brazil and Guyana (Ule 8372, Graham 105) as well as from Piauí State, Brazil (Menezes 1007, Pires 17348) the indumentum is sparse.Some are intermediate between this pattern and the pattern 2: they have leaves with very short, almost nonexistent, trichomes (e.g.Magalhães 8627, from Anápolis, Goiás, Brazil, and Trujillo 10780, from the Parguaza river in the State of Bolívar, Venezuela).Menezes 1007, because of its robust stem, resembles plants of pattern 8. Steyermark 113210, from K, fit within this pattern, while those from F resemble pattern 1.The same is true for Maguire 31679 (sheets 1 and 2, respectively).Philipson 2320 (type collection of V. macarenensis) belongs to this pattern, although it is more robust and, not considering the indumentum, it is very much the same as the plants from the type collection of V. dumitiana, within pattern 6. Hatschbach 60310, in light of the delicate stem, resembles the plants of pattern 3. The palisade parenchyma is inconspicuous in Rosa 2018 (figure1D), as in some collections of patterns 1 and 8.
Dawson 14674, Irwin 12454, Pirani 1683)and the Cachoeira da Misericórdia along the Juruena river in Mato Grosso (Rosa 1994), both in Brazil, as well as at Pakaraima mountains (Maguire 32307) and the Mazaruni river (Tillet 43855) in British Guyana.The plants from Rivadavia Lopes 181, from Chapada dos Veadeiros, are very similar to plants of pattern 3. Mato Grosso, Brazil (Rosa 2018), and from Serra da Mesa in Minaçu (Mello-Silva 548) and Caiapônia (Anderson 12421), both in Goiás, Brazil, all possess very long and dense, sometimes extremely hirsute, trichomes.The collections from Serra do Aracá (

Table 2 .
Distrito Federal, and in the Chapada dos Veadeiros mountains between Guaraí, to the north, and Caldas Novas, to the south.Disjunct populations are found in the Chapada Diamantina in Bahia State, in the localities of Abaíra (Ganev 623) and Piatã (Mello-Silva CFCR7386); in Cachimbo farm (Cordeiro 1191) and Serra do Cachimbo (Nascimento 540) in Mato Grosso; and in Patrocínio (Irwin 25614) in Minas Gerais.In Bahia, the indumentum is extremely dense and woolly.Plowman 8291, from Tocantins State, shows leaf sheaths well spaced one from another, giving the plant an unique aspect.Hatschbach 58358, Ratter 45252, and Splett 277 possess all of the characteristics of this pattern, although the dry leaves are involute, typical of plants of pattern 8. Ule 3142 (type specimen of V. macrosiphonia) could not be located, but to judge by the locality where it was collected, as well as from its description, it must belong to this pattern.figure8C).Includes the types of V. dumitiana and V. maudeana.The plants are notable for their almost glabrous leaves, sometimes ciliated only at base.They occur in Colombia in the cerro Isibukuri; in Venezuela, in cerro Duida (e.g.Maguire 29156) and nearby along the Cunucunuma river (e.g.Steyermark 125834); in Brazil, they occur in Serra do Aracá (Pires 15053), and along the upper stretches of the Mazaruni river in Guyana (e.g.Boyan 7).Collections of pattern 8 of Vellozia tubiflora, showing variation on flower colour and dimensions of individuals.
Semir 20642, Menezes 1004), which have a slightly heavier indumentum, are intermediate between this pattern and pattern 5.The foliar anatomy of plants from Serra do Cachimbo is similar to the collection Rosa 2018, of the pattern 4.These differ from the remaining in that the fibre bundles on the adaxial leaf surface are exiguous or absent, and they have a large quantity of stomata on the adaxial surface.Rizzo 7242, of unknown origin, possesses all the characteristics of the plants from Cristalina, although their leaves are involute.