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A Revision of the Genus Polylepis (Rosaceae

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Polylepis multijuga PiIger

Polylepis pauta Hieronymus ,

Leaves slightly obtuse at the ends of the branches, imparipinate with 4–9 pairs of leaflets (Figure 14a), obtuse in outline, 2.4–6.6 cm wide and 3.4–11.7 cm long; rachis lanose; point of leaflet attachment hispid on the upper surface, the long (to 3 rnm) trichomes (Figure 14c) usually mixed with multicellular, glandular trichomes and resinous exudate; stipular sheaths reddish brown, sericeous on the outside. Leaflets (Figure 14b) ovate, acuminate at apex, oblong or lanceolate in outline, second pair from terminal leaf usually the largest, one of this pair 0.6–1.4 cm wide and 0.8–3.3 cm long; edges jagged; apex acute; base odd cordial; upper leaf surface smooth or slightly tomentose, especially in the midvein depression; lower surface serriceous with the white or gold trichomes entirely confined to, or most conspicuous on, the veins. Flowers (Figure 14e) perfect except for possibly some plants from Peru (Cuzco) which appear to be male sterile, 0.3–0.6 cm in diameter; sepals 3 or 4, ovate, green, outer surface serricous; stamens 5-12, red, sericeous; stigma expanded, style hispid at base.

Fruit (Figure 14f) irregularly shaped, rhomboid to cylindrical with numerous, unequal, smooth to villous, thin spines, 0.3–1.4 cm wide including spines, 0.3–1.2 cm long. This species appears to grow in relatively inesian habitats, usually mixed with or at the upper edge of the montane forest. Specimens have been collected as low as 1800 m, the lowest altitude recorded for any species of the genus within.

Orthologically, Polylepts pauta is in many characters intermediate between P. rnultijuga and P. . lariugtnosa on one side and P. sericea on the other. It differs from the previous two taxa b) by smaller leaves and inflorescences, reduced hairiness on the leaf sheaths and lobes and on the inflorescences and restriction of hairiness on the lower leaf surfaces to the veins.

Polylepis sericea Weddell

However, examination of a large number of specimens from across the range shows that plants similar to a form characteristic of one area often occur in a population from another part of the range. In fact, the amount, length and color of the pubescence on the lower surface of the leaflets varies greatly throughout the series. In Colombia, the trichomes of the lower leaf surfaces tend to be slightly shorter than those most common in Venezuela, while plants from C o r a o h and in Peru plants from Caraz in Ecuador have pronounced pubescence on the upper leaflet. surfaces as well, similar to that of P.

There is also some variation in the shape of the tip of the stipule sheaths. Leaves at the ends of branches condensed, oddly pinnate with 1 pair of leaflets; trullate in outline, 1.7-3.3 cm wide, 3.1-3.4 cm long; rachises pubescent with scattered multicellular glandular trichomes; point of attachment of leaflets (Fig. 18b) with a small tuft of long straight unicellular trichomes; stipular sheaths declinating away from the peduncle, a compressed lobe on the upper part and outer side. REPRESENTATIVE COLLECTIONS EXAMINED.-The only collections seen are those listed as the type species and its synonym (Fig. 19).

However, as Macbride points out, the dense, straight trichomes covering the undersides of the leaves are more similar to those of P populations. Due to the coarse, compact covering of straight trichomes on the leaflets and the single pair of leaflets, this taxon is easily distinguished from other members of the genus.

FIGURE  I6.-PoZyZepis  sericea:  a,  branch  ( X   0.5);  b ,   lower  leaflet  surface  ( x   3);  c,  point  of  leaflet  attachment, lower  surface  ( X   4);  d, flower  ( x   6 ) ;   e,  fruit  ( x   6)
FIGURE I6.-PoZyZepis sericea: a, branch ( X 0.5); b , lower leaflet surface ( x 3); c, point of leaflet attachment, lower surface ( X 4); d, flower ( x 6 ) ; e, fruit ( x 6)

Polylepis pepei, new species

The tendency for the leaflet apex to appear three-toothed due to projection of the midvein is also unique in the genus. Flowers (Figure 240 perfect, 0.4-0.6 cm in diameter; sepals four, ovate, sericeo-lanose on outer surface; stamens 6-16, purple, locules cracked to glabrous, base of style fleshy. The two may are most easily distinguished on the basis of the plumage of the lower leaf surfaces and the stipule sheaths.

In all specimens of the species examined, at least a small group of glandular trichomes exists at the point of attachment of the leaflet. Leaves compressed at the ends of branches, saturated with 1-3 pairs of leaflets (Figures 28a, 29a); outline rhombic or obtrullate, 1.4-4.9 cm wide, 1.8-7.9 cm long, rachises densely lanate mixed with glandular trichomes;. Flowers (Figures 28d, 29c) perfect, 0.9-1.0 cm in diameter; sepals 3-4, ovate, slightly lanate on the outer surface and on the tips of the inner surface, the outer surface often with variable amounts of glandular trichomes;.

1In a population, the greatest variation is in the size of the leaflets and the density of the trichomes on the underleaf surfaces. The dense, tomentose (pannose) covering of the underside of the leaflets of these two taxa is never found in P. incana. This variability has unfortunately caused a taxonomic confusion in the treatment of the species.

Establishment of Polylepis sericea seedlings in the Paramo Zone (.%lpine) of the Venezuelan Andes.

FIGURE  20.-Polylepis  pepei:  a,  branch  ( X   0.5);  b,  leaf,  lower  surface  ( x   3);  c,  point  of  leaflet  attachment, upper  surface  ( X   12.5);  d,  flower  (T%ith stamens  missing)  ( X   4.5);  e,  fruit  ( X   4.5)
FIGURE 20.-Polylepis pepei: a, branch ( X 0.5); b, leaf, lower surface ( x 3); c, point of leaflet attachment, upper surface ( X 12.5); d, flower (T%ith stamens missing) ( X 4.5); e, fruit ( X 4.5)

Polylepis reticulata Hieronymus

Polylepis weberbaueri Pilger

Leaves very compressed at the ends of the branches, sometimes giving the appearance of a wrinkled arrangement (Fig. 24a), oddly pinnate with 3-5 pairs of leaflets, bordered in outline, 0.9-2.4 cin wide and 1.5-5.0 cm long; rachises denszly lanate or lanate with dark red glandular trichomes; attachment point of leaflet (Fig. 24e) with tuft of matted multicellular glandular trichomes; leaf sheaths with short blunt spurs, outer surface glabrous with long straight silky trichomes projecting from the lower surface. Leaflets (Fig. 2 4 b) oval to inverted ovate in outline, first (rarely second) pair from apex. Inflorescence pendulous, 2.1-3.4 cm long, with 3-7 flowers collected at the terminal end; petals small, 0.1-0.2 cm long, lanate or scaly; rachises lanose.

Fruit globose in outline, irregularly tilted, the spines flat, body of the fruit and spiders short; about 0.29-0.57 cm wide including spines cm long. DISTRInu?.Ion.-Central Ecuador in the province of Pinchincha south to north of Peru in the Department of Ancash, mainly on dry slopes of the Andes (Figure 25). DISCUSSION.-In many respects Polylepis weberbaueri appears to be the ecological equivalent in the drier Ecuadorian Cordillera of P.

In our collections in southern Ecuador (Simpson 8532) on the slopes of mountains near Canar, we found plants with mature, lower leaves characteristic of P. H e recognized (and listed) intermediate forms between the pure lanose type and the lanose gland type.

FIGURE  25.-Collection  localities of  Pol)lepis  weberbaueri.
FIGURE 25.-Collection localities of Pol)lepis weberbaueri.

Polylepis quadrijuga Bitter

Since a grading of glandular trichome density exists within the populations from Chimborazo, formal recognition of a separate variety seems unnecessaryj.

FIGURE  26.--PolyZepis  qzradrijuga:  a,  branch  ( x   0.5):  b,  undersurface  of  leaflets  ( x   3):  r ,   point  of  leaflet attachment, upper surface  ( x   6);  d,  e ,  flowers  ( x   6);  f ,   fruit  ( x   6)
FIGURE 26.--PolyZepis qzradrijuga: a, branch ( x 0.5): b, undersurface of leaflets ( x 3): r , point of leaflet attachment, upper surface ( x 6); d, e , flowers ( x 6); f , fruit ( x 6)

Polylepis besseri Hieronymus

Inflorescence pendulous (or more or less straight, if very short cm long, with 3-12 flowers; flower bracts lanceolate, glandular on the outer surface; rachis glabrescent with scattered glandular trichomes or slightly woolly. Guerraloma, 2800 m, May 1958, Carderias 572f (USA) DISCUSSION.-1. Polylepis besseri undoubtedly consists of the most polymorphic and confusing group of populations of any species of the genus. The same confusion that I experienced in the treatment of these populations is evident in treatments of previous workers.

In the original description of the species of Hieronyinus, several collections were cited from different parts of Bolivia. Bitter specifically stated that Hieronymus' species was an unnatural species containing more than one species, and he separated some of the collections out at P. In addition, Bitter described several other species on the basis of one or two collections.

Assigning different populations to a series of species as proposed by Bitter does not make sense in light of the number of intermediate populations and variation within populations. Among populations, the variation is in the amount of leaf margin formation or curling, the amount of roughness of the upper leaf surface, the extent of glandular trichomes on the veins of the lower leaf surfaces, and the amount of fusion of the spines. of fruit to form spiny wings.

FIGURE  28.--Polykpis  besseri  from  Colomi,  Bolivia:  a,  branch  ( X   0.5):  b,  point  of  leaflet  attach-  ment, upper  surface  ( x   3);  c,  point  of  leaflet  attachment,  lower  surface  ( X   3);  d,  flower  ( X   3);
FIGURE 28.--Polykpis besseri from Colomi, Bolivia: a, branch ( X 0.5): b, point of leaflet attach- ment, upper surface ( x 3); c, point of leaflet attachment, lower surface ( X 3); d, flower ( X 3);

Polylepis tomentella Weddell

DISCUSSION.—Included here in Polylepis tomentella are essentially all the forms originally included in the taxon by Weddell. Especially in central Bolivia, the pubescence on the underside of the leaves becomes less dense. It is possible that the two taxa approach each other morphologically due to the similarity of the habitats in which they occur.

The morphological characters that best separate the two, as discussed here, are the pubescence of the surfaces of the lower leaflets and the shape of the leaflet. Bitter never mentioned Bonpland's type of the species among any of his subspecific taxa, and so it seems possible that he was in fact unsure of the correct characterization of the name. As Macbride rightly pointed out, many of the varieties and subspecies are elements that belong within P.

DISTRISUTLON.-Northern Peru in the province of La Libertad and in the wet valley heads on the western slopes of the Peruvian Cordillera south to Lima and in central Peru from Huanuco through the Pampa de J u n i n to Huancayo (Figure 37). Polylepis incnna tends to be a species of drier regions, growing from Ecuador south to the eastern side of the Western Cordillera of Peru. Instead, after describing the species, he gave a general range of distribution and went on to enumerate eight varieties.

However, as with the other varieties of the species described by Bitter, there seems to be no justification for maintaining this variety or its subvarieties.

FIGURE  32.-PoZyZepis  tomentella  from  Cochabamba,  Bolivia:  a,  branch  ( x   0.5);  b,  cluster  of  leaves  ( x   3);  c,  upper  leaflet  surface  ( x   12.5);  d, flower  ( x   6);  e,  fruit  ( x   6)
FIGURE 32.-PoZyZepis tomentella from Cochabamba, Bolivia: a, branch ( x 0.5); b, cluster of leaves ( x 3); c, upper leaflet surface ( x 12.5); d, flower ( x 6); e, fruit ( x 6)

Polylepis incana Humboldt, Bonpland and Kunth

Polylepis racernosa Ruiz and Pavon

Polylepis australis Bitter . ,

Leaflets broadly lanceolate, irregularly ovate, the first or second pair from the terminal leaflet the largest, one of the pair 0.3-1.5 cm wide, 1.0-4.0 cm long; Inflorescence drooping, 1.8-7.3 cm long, with 2-12 flowers, bracts lanceolate, scaly, rarely glandular, 3-6 mm long, stipitate with spreading. Flowers (Fig. 38c) perfect, 0.7-1.0 cm in diameter, sepals 3 or 4, ovate, outer surface glandular or hairy, stamens 8-16, anthers with tuft of straight trichomes at apex;

In general, the fruits of this species are more winged than those of any other in the genus. Further complications in the taxonomy of the species arise because Bitter used different specimens from the same collections as types of some of his varieties (e.g. Lorentz 310, Hieronymus s.n.). As the actual specimens used by Bitter were destroyed in Berlin, and as he evidently considered collecLioris to be polytypic, it is impossible to refer duplicates of these collections to his various varieties.

Although the variety descriptions might give some indication of his labeling of individual specimens, there is so much variation that there is no labeling of parts of the collections. There are no specimens associated in the variety description; however, there are extant isotype collections that he listed for various subgenres.

FIGURE  38.--PoZyZepis  australis:  a,  branch  ( x   0.5);  h,  point  of  leaflet  attachment,  upper  surface  ( x   3):  c,  flower  ( x   6);  d, fruit  ( x   6)
FIGURE 38.--PoZyZepis australis: a, branch ( x 0.5); h, point of leaflet attachment, upper surface ( x 3): c, flower ( x 6); d, fruit ( x 6)

Gambar

FIGURE  2.-Branching  and  flowering  patterns  of  Polylepis  species:  a,  congestion  of  leaves  of  P
FIGURE  3.-Leaflet  surfaces  of  Polylepis  species:  a,  underleaf  surface  of  P
FIGURE  5.-Cross  sections  of  the  leaflets  of  Polylepis  reticulntn  ( x   105), showing  the  intraspecific  variability  in  the  number  of  cell  layers  of  the  epidermis  and  the  thick  cuticle  characteristic  of  members  of  the  w t i c u
FIGURE  B.-Cross  sections  of  Polylepis  besseii  leaflets  under  regular  ( n )  and  polarized  ( b )   light  show  the  in\aginated  lower  epidermis,  the  small  cell  size,  and  the  packed  arrangement  of  the  cells  characteristic  of  membe
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