Roupala montana var. paraensis (Sleumer) K.S.Edwards

  • Authority

    Prance, Ghillean T., et al. 2007. Proteaceae. Fl. Neotrop. Monogr. 100: 1-218. (Published by NYBG Press)

  • Family

    Proteaceae

  • Scientific Name

    Roupala montana var. paraensis (Sleumer) K.S.Edwards

  • Type

    Type. Brazil. Para: Peixe Boi, 21 Jul 1907 (fl), Siqueira s.n. (= R. S. Rodrigues). MG 8263 (holotype, G, photo 27823 F, GH, MO; isotypes, BM, C, MG, fragm. LIL, RB, S, n.v., U).

  • Synonyms

    Roupala nervosa Klotzsch, Roupala lucens Meisn., Roupala brasiliensis var. subintegra Meisn., Roupala glabrata Klotzsch, Roupala longipetiolata Pohl, Roupala rhombifolia Mart., Roupala tristis Mez, Roupala organensis Gardner, Roupala rhombifolia var. paraensis Sleumer

  • Description

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    Variety Description - Leaves drying dark olive-green, narrowly elliptic to elliptic, length:breadth ratio 2-3:1, soon glabrous beneath; indumentum, if present, of short hairs only, if persistent, ferruginous red-brown; base narrowly cuneate. Inflorescence 2-13 cm long, indumentum dense to sparse, red to dark brown, the hairs straight, appressed. Flowers not densely clustered at anthesis, drying dark brown. Ovary indumentum appressed, hairs extending 1-2 from base. Field characters. Shrub or small tree 2-10 m tall, 6-11.5 cm diam. Branches ascending, crown dense. Bark gray to grayish-brown; external bark 3 mm thick, brown; internal bark 8 mm thick, pink, bitter to taste. Wood yellow to cream, or cream to red; hard. Leaves coriaceous, dark green, shiny above, new leaves ferruginous beneath. Inflorescence pale brown to dark brown to ferruginous. Buds green tinged orange-brown to yellowish-brown. Flowers fragrant, white, cream, pale yellow, greenish-white, or yellowish green; stigma pale green; ovary ferruginous to brown; nectary white. Fruit green, lightly ferruginous, maturing brown to grayish-brown with persistent style. Seeds pale brown.

  • Discussion

    Roupala montana Aubl. is a widespread ochlospecies (White, 1962; Prance, 1972; Cronk, 1998). Small variations between specimens have invariably lead to a proliferation of names, but an examination of over 900 specimens, together with extensive observations in the field, has resulted in much synonymization. A broad species concept has been adopted here to address the variability, with recognition of four varieties that represent extremes of that variation.

    If one regards the specimens from Central and northern South America, Roupala montana is easily defined. It occurs as a shrub or tree, in savanna or forest, from sea level to high altitudes. The leaves are principally ovate, widely ovate, or very widely ovate in shape, rounded to cordate at the base, obtuse then attenuate at the apex, the margin more often entire or entire and undulate rather than serrate or crenate, glabrous when mature, and usually drying glaucous green. Inflorescences, pedicels, and perianths generally have a short-pilose indumentum, hairs light brown and straight, while ovary hairs are off-white through dull yellow to light brown and short. This suite of characters constitutes R. montana var. montana.

    However, moving south into Bolivia, Paraguay, and particularly southern Brazil, the species becomes more variable. Leaves demonstrate a considerable variety of form: narrowly to widely elliptic, oblong to widely oblong, lanceolate, and narrow to widely obovate, as well as narrow to wide ovate, which predominates in the north. Leaf apices and bases are correspondingly variable. Leaf margins are more often serrate than entire, with indumentum often persisting, especially on the underside of the leaves. Leaves dry green-brown or ochre to mid or even dark brown, more often than glaucous. Inflorescences, pedicels, and perianths are covered by a tomentose, orange-brown indumentum. The ovary indumentum is very dense, consisting of long hairs. In the southern range of this species most of the material was formerly placed in Roupala brasiliensis which is here relegated to varietal level because of the overlapping polymorphic variation. Like the type variety var. brasiliensis is represented by shrubs and trees occurring in a similar variety of habitats. Histograms representing the number of individual specimens of Roupala montana var. montana and var. brasiliensis against variations in lamina length, lamina breadth, petiole length, number of lateral vein pairs, leaf shape, leaf base, and leaf apex shapes are presented in Figure 35. Sixty specimens of each variety were scored to produce these histograms. Roupala montana var. brasiliensis resulted in better-defined peaks for the graphs of lamina length and breadth than var. montana, which showed a greater distribution of individuals between the size classes. Petiole lengths of the two species displayed almost identical distributions across the size classes.

    Roupala montana var. brasiliensis is much more variable with respect to leaf shape than var. montana. It is represented by five additional categories and has quite an even distribution in the number of individuals among most of the categories. In Roupala mon- tana var. montana almost half of the specimens (45%) are ovate, while 83% are contained between the extremes of narrow and wide ovate. The leaf apex histogram (Fig. 35C) showed much similarity between var. montana and var. brasiliensis, the majority of both specimens being either acute and attenuate or obtuse and attenuate. Leaf bases of var. montana are more frequently obtuse (47%) than acute (32%), while the reverse is true of var. brasiliensis (22% versus 53%, respectively) for the 60 specimens of each examined.

    Interestingly, the considerably greater variation in leaf shape for Roupala montana var. brasiliensis does not produce greater variation in the number of lateral vein pairs (Fig. 35F). In var. montana, 95% of specimens have between 4 and 8 pairs, while in var. brasiliensis, 83% of specimens have only 5-7 pairs.

    Of the 60 specimens of Roupala montana var. montana, 46 had entire leaf margins, while only 14 were serrate or crenate. The figures for R. montana var. brasiliensis are 13 and 47, respectively, pointing to a possible relationship between type of leaf margin and indumentum characteristics. However, because both are apparently continuous variables, they cannot be weighted too heavily, although the trends are noted.

    There is, therefore, no convincing evidence from vegetative characteristics which could be used to separate Roupala montana var. brasiliensis from R. montana var. montana at a level other than varietal. With regard to fertile characters, 20 specimens of each of the two groups were examined, and measurements were made of common bract length and breadth (at midlength along the inflorescence) and the lengths of pedicels, flowers, nectary lobes, and anthers.

    Common bracts of Roupala montana var. brasiliensis tend to be broader and longer than those of R. montana var. montana, 0.4-1.2(1.5) mm broad and (0.5-) 1-2.5 mm long compared with 0.3-0.8 mm and (0.3-)0.4-l mm, respectively. Pedicel, flower, nectary lobe and anther lengths show equal variation between the two varieties (1-) 1.5-3(-4.5) mm, (5-)7-9(-10.5) mm, 0.2-0.6 mm and (1-) 1.5-2.8 mm for successive characters of var. brasiliensis and 1.5-3(-4.5) mm, (5-)7-8.5(-10) mm, 0.3-0.5 mm and (1-) 1.5-3 mm for var. montana.

    The evidence for separation of these varieties on floral characters is no more convincing than that for vegetative characters, the most outstanding difference being in the measurements of the common bracts. However, intermediate specimens between the varieties showed intermediate lengths of common bracts. The shape and size of common bracts alone are unlikely to be at the root of any evolutionary mechanism which may have caused possible speciation.

    Fruit size and shape within this species is very variable and even in association with vegetative and floral characters, provides no further basis for separating the varieties into two species. The fruit is an awkward organ to use when looking for characters in herbarium specimens because the follicles tend to open as the specimen dries, even when obviously immature. Thus, one can never be certain whether morphometric and morphological differences reflect true taxonomic differences rather than merely different stages of ontogeny. (Three specimens in Paraguay have more beaked fruit apices than is usual in South and Central America, two of which have a persistent sandy-colored tomentum). Whether a future increase in collections consisting of specimens with both flowers and fruit together (from the same individuals) will illuminate useful fruit characters brought to attention through the more conspicuous floral differences, is yet to be seen.

    A final attempt to look for character correlations was based on Anderson’s (1952) concept of introgressive hybridization, with evidence provided by pictorial scatter diagrams. A random collection was made of 20 specimens each from Roupala montana var. montana, var. brasiliensis, and their intermediates. The two morphometric variables of leaf length and leaf breadth were selected and the measurements recorded using the average mature leaf size on each specimen. Roupala montana var. brasiliensis was expected to have a longer, narrower leaf than var. montana. Four qualitative characters that clearly show variation across the two varieties and their intermediates were chosen in order to see whether any of them were linked with the leaf measurement data. These qualitative characters were leaf margin type (entire, intermediate [undulate/with slight presence of teeth], or toothed), and leaf, perianth, and ovary indumentum (weak, intermediate, or dense). The analysis was repeated using additional morphometric variables: (1) the ratio of flower length to pedicel length, and (2) the ratio of leaf length/petiole length to flower length/pedicel length.

    The diagrams showed that no correlations were evident in either the two sets of morphometric data or between any of the qualitative and morphometric data. With larger data sets, perhaps some tendencies towards character correlation would start to appear, but the lack of any positive evidence from this pilot study discouraged a more extensive survey. The distribution of characters suggests reticulate variation, with no evident linking of characters into complexes. No trends were evident to differentiate the varieties on the basis of habit, altitude, habitat, or geographical area. Roupala montana var. brasiliensis is a shrub l-2 m tall, a small tree, or a tree to 25 m tall found at 200-1800 m alt. in primary and secondary forest and in the open habitats of cerrado, capão, campo sujo, capoeira, and gallery forest beside these habitats. Roupala montana var. montana has the same habit and occurs in the same habitats, between sea level and 1600 m. Distribution maps of the two varieties and intermediates (Figs. 33, 34A) show that var. brasiliensis and var. montana are sympatric, although the range of var. montana is much more extensive. The intermediates always occur where there are populations of both groups.

    The result of this data sifting is that no positive evidence has been found for maintaining Roupala montana var. brasiliensis apart from var. montana at anything above the status of variety. While they are clearly separable in their extreme forms, their sympatricity, the actual characters and degree by which they differ, and the presence of intermediate specimens together make it impossible to accept them as different species.

    With this apparent reticulate distribution of characters among specimens of Roupala montana, it is possible that the extreme forms were once distinct species occupying the same habitats in the same geographical region in the southern part of the continent. Perhaps the two species began to hybridize introgressively, generating progeny that carried mixtures of characters of either parent, which then hybridized further with the pure parental types and other hybrid offspring. In this way, character complexes originally associated with each species may have broken down causing complete intergrading between the original taxa.

    Another theory, of course, is that two distinct taxa never existed as such, and that one highly variable species has resulted from a phenotype responding very strongly to its environment. A species which has the ability to produce a large amount of wind-dispersed seed is likely to be a good pioneer species, readily adaptable to new habitats and conditions. Johnson & Briggs (1975) postulated that the tropical species of Proteaceae (Grevillioideae) originated as trees in the forest situation, and from there became invasive and often dominant in open habitats as seed was carried out.

    A greater degree of separation is evident between Roupala montana var. montana and the varieties paraensis and impressiuscula than between R. montana var. montana and var. brasiliensis, with almost no intermediates. However, the main distinguishing character of indumentum color is not deemed sufficient to recognize var. paraensis and impressiuscula at the specific level, especially since the indumentum color itself is not consistently dark red, but may grade to a lighter orange. While R. montana var. montana (and var. brasiliensis) has a golden, pale brown, orange-brown or mid brown indumentum on rachises, perianths, ovaries, and the underside of leaves, vars. paraensis and impressiuscula have distinctly red-brown hairs. The latter group usually have an angular leaf base which is broadly to narrowly cuneate, as compared with acute to obtuse to rounded. While there are other subtle differences also highlighted at couplet 2 in the key, most morphometric variables of components in vars. paraensis and impressiuscula fall within the ranges of those components in R. montana var. montana, making it unrealistic to recognize these varieties at specific level.

    The scatter diagrams of the correlation analysis showed the distribution of four character states within Brazilian Roupala montana: (1) appressed versus tomentose inflorescence indumentum, (2) red-brown indumentum v. golden, pale brown, orange-brown or mid brown, (3) cuneate leaf base v. acute/obtuse/rounded, and (4) glabrous undersides of leaves v. persistent indumentum. The most striking information shown in the map is that red-brown indumentum is restricted to a high altitude, narrow coastal strip of SE to NE Brazil. These plants generally also have a cuneate leaf base, though not always, and specimens of the other indumentum color cate- gory, both with tomentose and appressed indumentum, also at times have cuneate leaf bases. The red-brown indumentum character is found in a band from Paraná to Pará, while the contrasting indumentum color is widespread across all of Brazil, including this band. The tomentose inflorescence indumentum character and persistence of indumentum on the undersides of leaves would appear to be closely correlated, particularly in southern Brazil and Bolivia, although leaf indumentum is nonpersistent on scattered specimens with tomentose inflorescences from NE Brazil to Pari Cuneate leaf bases are apparent in a broad arc from southern Bolivia through southern Brazil to NE Brazil and Pará, as compared with acute/obtuse/rounded leaf bases, which are widespread in all of the country.

    It seems most appropriate to maintain the taxa Roupala paraensis and R. impressiuscula as varieties of R. montana rather than as species since they are based on such slight character differentiation and in parts of their range are hard to differentiate in the field. The taxonomic classification proposed here re-mains unsatisfactory and needs further field work to resolve.

    Distribution and Ecology: Occurs in the coastal range of the Serra do Mar of Santa Catarina, Sao Paulo and Rio de Janeiro, Brazil, and farther inland in the Serra Paranapiaçaba of Paraná, the Serra da Mantiqueira of Minas Gerais and São Paulo, the Serra do Espinhaço, and also in Minas Gerais, Para, and Tocantins. There is one puzzling disjunct at Peixe Boi, several degrees north in Para. The altitude range of the species is generally between 1200-2000 m, although it is characteristically at lower altitudes in Santa Catarina, 350-1000 m, and it reaches 2300 m on the Serra do Caparão, Minas Gerais. A plant of campo rupestre, high altitude campo, cerrado, open campo, and capão, often among rocks, it also extends into small pockets of forest and strips of gallery forest.

  • Common Names

    Carne de vaca, carvalho, congonha, congonha vermelha

  • Distribution

    São Paulo Brazil South America| Tocantins Brazil South America| Bahia Brazil South America| Rio de Janeiro Brazil South America| Santa Catarina Brazil South America| Minas Gerais Brazil South America| Paraná Brazil South America|