BotanualJournal of the Linnean So&& (1999), 129: 315-332. With 4 figures @
Article ID: bojl. 1998.0223, available online at http://www.idealibrary.corn on I D E L L CI
Revision of the genus Fascicularia Mez (Bromeliaceae) GEORG ZIZKA'*, RALF HORRES', E. CHARLES NELSON' AND KURT WEISING3 I Botany /Paleobotany, Forschungsinstitut Senckenberg and Universip of Frankjirt, Senckenbqanlage 25, 0-60325Frankjirt/Main, Germany. 'lippitiwitchet Cottage, Hall Road, Outwell, Wubech PE14 8PE. 'Plant Molecular Biology, Biozentmm, JV200, Uniuersip of Frankjirt, Marie-Curie-Str. 9,0-60439 Frankjirt/Main, Gennany
Received Mgy 1998; accepted for publication September 1 9 9 8
The genus Fmcicularia Mez is revised as part of a study of the Bromeliaceae for the Flora de Chile. Morphological and anatomical investigation of herbarium and living material from cultivation as well as DNA-studies (RAPDs) in cultivated material has led us to conclude that Fmcicularia bicolor (Ruiz & Pav.) Mez, the only one species in the genus, comprises two subspecies which are distinguished by their leaf anatomy and morphology. 0 1999 The Linnean Society of London
ADDITIONAL KEY WORDS:--Chile - morphology - RAPDs.
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classification - Fmcicularia bicolor - molecular data
CONTENTS
Introduction . . . . . . . . . . . . . Results . . . . . . . . . . . . . . Morphology . . . . . . . . . . . Anatomy . . . . . . . . . . . . Geographical distribution and ecology . . DNA analysis . . . . . . . . . . . Descriptions and key . . . . . . . Acknowledgements . . . . . . . . . . References . . . . . . . . . . . . . Appendix: specimens seen . . . . . . .
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315 316 316 317 318 320 322 330 330 332
INTRODUCTION
Fascicularia as represented here is monotypic. The only species, Fascicularia bicolor (Ruiz & Pav.) Mez, was first described as Bmmelia bicolor by Ruiz & Pavon (1802).
* Corresponding author. Email:
[email protected] 002+4074/99/040315+ 18 $30.00/0
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Vegetatively, Fascicularia is similar to Ochagauia Phil. Both genera are endemic to central and southern Chile and comprise a group of terrestrial or epiphytic bromeliad species with more or less succulent leaves. Following Ruiz & Pavon, Molina (18 10) and Gay (1853) recognized Bmmelia bicolor as the only species of this group. A few years later Philippi (1856) described the genus Ochagavia, at first including only one species endemic to Isla Robinson Crusoe, Juan Fernhndez Islands (Ochagauia elegans Phil.). Shortly afterwards, Philippi (1857-1858) defined another genus, Rhodostachys, with a single species, R. andina Phil. Delimitation of the genera was not clear and this led to a number of new combinations by Bentham (in Bentham & Hooker, 1883) and Baker (1889). The taxonomy and nomenclature of this group of genera were further complicated by new names and combinations published in the late 19th century based on cultivated material from European gardens without preservation of type specimens. In a footnote to his treatment of the Bromeliaceae for Martius’ Flora Brasiliensis, Mez (1 894) created the new genus Fascicularia, including three species formerly placed in Rhodostachys ( E bicolor, R litoralis, E pitcairniijilia) and separated them from the remaining species of Rhodostachys on the basis of petal appendages and pollen morphology. Mez (1896) realized that Ochagavia elegans and the species of Rhodostachys were closely related and therefore united the two genera, at first erroneously selecting the younger name Rhodostachys. This was corrected by Mez (1934) and Smith & Looser (1934), who published the presently accepted concept for the species group, comprising the genera Fascicularia and Ochagavia. Ochagavia can best be distinguished from Fascicularia by the relative lengths of the style and the stamens, by sepal shape, petal appendages, petal colour, and pollen morphology (see also below). Although flowering plants of Fascicularia and Ochagavia display striking differences, the relevant characters are difficult to study in herbarium specimens. Moreover, only some of the available specimens include floral parts (in the present study 39/89); even then, because of the mode of anthesis in bromeliads, only a single or a few fully developed flowers may be available for investigation. These genera therefore exemplify a major problem in bromeliad taxonomy which is that for most genera and species, the number of herbarium specimens available is comparatively small (due to difficulties in accessing, drying and handling of these plants) and thus our knowledge of variability of characters regarded of taxonomic importance is insufficient. Modern revisions including characters beyond morphology are scarce. Recently, comparative studies of morphology and micromorphology (e.g. Bohme, 1988; GroR, 1988; Schill, Dannenbaum & Jentzsch, 1988; Brown & Gilmartin, 1989; Brown & Terry, 1992), palynology (Halbritter, 1992), anatomy (Horres & Zizka, 1995) and DNA sequences (Clark et al., 1993; Ranker et al., 1990; Linder & Kellogg, 1995; Terry & Brown, 1996; Terry, Brown & Olmstead, 1997) have improved our knowledge about suprageneric grouping and relationships of the family. RESULTS
Morphology For the delimitation of species, Mez (1896, 1934), as well as Smith &Downs (1979) who adopted Mez’ classification, relied principally on the following morphological
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Figure 1. Scatter plot of width leaf blade us width leaf sheath measured from 47 herbarium specimens of Fnscicularia bicolor ssp. bicolor (0) and canaliculata (4.
characters: length of floral bracts relative to length of flower, flower length, as well as shape and apex of the sepals. Our morphological studies concentrated on the significant characters of leaves, inflorescence and flowers. For the 89 available herbarium specimens (including duplicates; see also Appendix), the sizes (length/ width) of leaf sheath and leaf blade were measured, and from 39 specimens measurements of floral parts were obtained. Our investigations revealed that flower and bract length are highly correlated, the bracts being always shorter than the developed flower. The absolute length of the flower appears to be highly variable (with a tendency toward longer flowers in ssp. bicolor), but does not seem to be of value in separating any groups within the genus. Although considerable variation in shape and indumentum occurs, all sepals were observed to be ovate to oblong, apically retuse or praemorse, with a short inconspicuous mucro. In our opinion, none of these characters serves to distinguish or characterize groups within Fmcicularia. Differences in shape and morphology of the leaf blade led us to define two taxa within Fmcicularia (see also below), although these cannot be clearly separated using other morphological characters (Nelson & Zizka, 1997). The width of the leaf blade is the only, easily accessible morphological character that can be obtained from herbarium specimens which correlates fairly well with the different leaf blade types (Fig. 1). Discrimination of the two taxa was also supported by the results of factor analysis and discriminant analysis (performed with software SPSS) based on 13 morphological characters (length/width of leaf sheath, leaf blade, inflorescence, floral bracts, sepals, petals as well as flower length) measured from 39 herbarium specimens.
Anatomy The anatomy of the leaf-blade in Fmcicularia bicolor is similar to other investigated Bromelioideae. The species has a one-celled sclerotic hypodermal cell-layer. The
inner hypodermal layers are sharply differentiated from the (sclerotic) outer layer and thc inner, assimilating tissue (chlorenchyma), and form characteristic water storage tissue. In Fascicularia, as in most other Bromelioideae but not in Tillandsioideae, water storage tissue is well developed only adaxially, and is usually lacking abaxially. Nevertheless, the amount of water storage tissue in the leaf-bladc (measured as percentage of area of transverse section) in R bicolor ssp. bicolor and Ochagavia carnea reaches the highest values among the investigated Bromclioideae, similar to Pitcairnioideae like Puya roe& or Hechtia glabra (Horres & Zizka, 1995). The two subspecies differ in leaf-shape and amount of adaxial water storagetissue. In ssp. bicolor, the adaxial surface of the leaf blade is not channelled longitudinally but is flat to slightly convex, In transverse section this can be easily seen, as well as the fact that thc water storage tissue is better developed than in ssp. canaliculata. In the latter subspecies, the leaves are conspicuously channelled longitudinally. The leaf blade in ssp. bicolor is considerably thicker than in ssp. canaliculata and usually wider (see fig. 1 in Nelson & Zizka, 1997). In herbarium material, determining the subspecies is difficult. Drying and pressing causes the water storage tissue to collapse and it cannot be fully restored. Comparison with herbarium leaves from cultivated material leads to only slightly different leaf-shape of thc two subspccies when the leaves are dried. Keeping the principal habitats of the two subspecies in mind (saxicolous near coast, epiphytic in forest), the differences in leaf anatomy might be regarded as mere variations corresponding to different ecological conditions without taxonomic value. However, the leaf characters proved to be constant in cultivation for a period of several years and correspond with two different banding patterns of RAPDs found in the cultivated material. Although not clearly separable by other morphological, especially floral, characters, the two groups are considered to be taxonomically distinct.
Geographical distribution and ecoloQ The specimens of Fascicularia bicolor we investigated were collected in Chile from app. 34"s to 42'24's (Fig. 2). Smith & Downs (1979) cite a specimen from 45'1 7's (Puerto Lagunas, Fricke s.n.; CM; not seen), which extends the distribution considerably further south, making Farcicularia the southernmost distributed member of Bromeliaceae. This area comprises mediterranean vegetation types in the north ('bosque esclerofilo'; Grau, 1995), temperate broadleaved forest ('bosque caducifolio templado') and the region of the Valdivian rain forest in the south. The area of the bromeliad genus Grekia extends nearly as far south, reaching Cucao on ChiloC (42"42'S, 74"05'W; Bettina Will, pers. comm.). Although there is no clear separation between the distributions of the subspecies, the natural range of ssp. bicolor extends farther north into areas with mediterranean climatc but including also the two above described more southern zones. Nevertheless, thc, ssp. bicolor is principally saxicolous, occurring in rocky habitats ncar thc coast. On thc other hand, E bicolor ssp. canaliculata appears to have its centre of distribution in thc area of temperate forests-the Valdivian rain forcst and the broadleaved tempcrate lorest--and usually grows as an epiphyte. These two forest types intergrade to some rxtcnt and share a considerable number of evergreen tree species (Schmithusen, 1956).
REVISION OF THE GENUS FASCICULARlA
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-35"
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-55" -76"
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-70" -68" Latitude S
'
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Figure 2. Distribution of Fascicularia bicolor ssp. bicolor (0) and canaliculata herbarium specimens (A) could not be determined as to subspecies.
-64"
(u)in Chile. A few
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Because of the difficulties in determining some herbarium specimens, a few specimens could not be assigned without doubt to one of the subspecies, and have been included in the map (Fig. 2) with a separate symbol. Further investigation of Fascicularia populations in Chile, especially careful description of leaf morphology when collecting herbarium specimens, will allow more precise information about the geographical distribution of the subspecies. Outside its natural distribution, Fascicularia bicolor has been introduced into gardens, and has become naturalized in western Europe. In favourable habitats, for example in the Isles of Scilly, western Ireland and the Channel Islands, Fascicularia colonies have spread, or been spread, to cover substantial areas of sand dunes and oak woodland.
DNA anabsis In recent years, the comparative analysis of DNA sequence variation has become increasingly important as a complementary tool to infer genetic relatedness among taxa (Bachmann, 1992; Hillis, Moritz & Mable, 1996). The random amplified polymorphic DNA (RAPD) marker technique described by Williams et al. (1990) is one of the most frequently used approaches of molecular taxonomy, mainly because of its technical simplicity. RAPD analysis makes use of the polymerase chain reaction (PCR; Mullis, FerrC & Gibbs, 1994) to generate polymorphic amplification products from anonymous regions of genomic DNA. Electrophoretic separation of RAPD products on agarose gels typically results in fingerprint-like banding patterns. Painvise genetic distance matrices can be calculated from the extent of band sharing, and the results analysed phenetically. In plants, genetic relationships based on RAPD data generally proved to correlate well with expectations from morphological, ecological or phenological data, especially if very closely related taxa were analysed (see e.g. Adams & Demeke, 1993; Ramser et al., 1996,1997; Furman et al., 1997). In the present work, we used the RAPD technique to assess intraspecific relationships among 12 Fascicularia bicolor accessions (eight of ssp. bicolor, and four of ssp. canaliculata). T o analyse the delimitation of the genus Fascicularia, we also included six accessions belonging to the related genera Ochagavia and Gei@a (i.e. four accessions of Ochagavia camea, and one accession each of 0. elegans and Grei@a sphacelata). All plant material originated from the living collection of the Palmengarten (Frankfurt/ Main, Germany). Genomic DNA was isolated from freshly harvested leaves by the method of Doyle & Doyle (1987), treated with FWAse, and used for PCR without further purification, RAPD reactions were carried out with arbitrarily chosen GCrich decamer primers (Operon, Alameda, FA), using the conditions described by Ramser et al. (1996, 1997). Of 14 primers initially tested, eight produced distinct polymorphic fingerprints and were used for further study. All assays were repeated independently, and only strong, reproducible bands were included in the final analysis. Banding patterns derived from primers OPG-7 and OPG-9 are exemplarily shown in Figure 3. Scoring for presence/absence of a band at a particular position resulted in 98 polymorphic characters across all primers (Table 1). This binary character matrix was transformed into a pairwise distance matrix using the Jaccard coefficient of similarity (Jaccard, 1908), and analysed by the unweighted pair group method using arithmetic averages (UPGMA; Sneath & Sokal, 1973). The resulting dendrogram is shown in Figure 4.
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Figure 3. RAPD analysis of Fasrirularia birolor and related genera using the Opcron primers GO7 (upper p a d ) and GO9 (lower panel). (a-h) R birolor ssp. birolor (eight accessions); (ikrn:) R birolor ssp. ranaliruluta (4 accessions); (11) 01,hagavia Plegans (1 accession), (o r) 0. c a n m (4accessions); (s) &pa sphacelala ( I accession). (C) negative control (no DNA template). Sample lanrs are flanked by molecular weight markcrs (lane M), positions of which are given in hasr pairs ( 1 00 Imse pair ladder, Gibco-BRL).
The 12 Fascicularia accessions are separated into two clusters which coincide well with the subspecies designation recognized on the basis of leaf blade anatomy and morphology. Thus, the RAPD data fully support the division of Fascicularia bicolor into two subspecies. Our DNA analysis also supports a clear-cut delimitation of Fascicularia from Ochagavia and Greigia. Somewhat surprisingly, however, the latter two genera are not well distinguished from each other. Whereas the four 0. carnea accessions form a separate cluster, the RAPD patterns suggest a higher similarity of 0. elegans (which is endemic to the Juan Fernaiidez Islands) to Greigia sphacelata than to the other Ochagavia accessions analysed. This finding is in some contradiction with morphological data, since Ochagavia species are morphologically very similar to each other, but clearly distinct from all Greigia species. The genera Ochuguvia and Greigia are currently under revision. The few DNA studies performed on Bromeliaceae so far (see Terry et al., 1997, and references cited therein) are mainly concerned with the still uncertain
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TABLE 1. List of decamer oligonucleotides used as random primers, their sequences, number of polymorphic products considered in DNA-analysis (only strong and reproducible signals considered) and fragment size range @p) Operon-Primer
number of polymorphic amplification products (size-range [bp])
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ITCGAGCCAG GAACCTGCGG AAAGCTGCGG ACTTCGCCAC GAACCTGCGG CTGACGTCAC AGGGCCCTCT ACTGGGACTC
G 07 G 09
G 10 G 15 Total
8 (480-1 700) 21 (420-1850) 9 (380- 1800) 13 (700-1800) I7 (750-2200) 12 (400-1850) 12 (300- 1800) 6 (950-2000) 98
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Figure 4. Phenogram constructed by UPGMA cluster analysis of a RAPD data matrix generated from (a-h) R bicolor ssp. bicolor, (i-m) R bicolor ssp. canaliculata, (n) Ochogavia elqans, (0-r) 0. carnea and (s) Greigia sphucelatu. Presencelabsence o f bands at 98 positions revealed by eight different Operon primers was transformed into a distance matrix using the Jaccard coefficient of similarity and analyzed by the UPGMA option of the NTSYS-pc software package (Row , 1993). The relative genetic distance is indicated by the horizontal bar.
suprageneric phylogeny of the family. Our RAPD results underline that there is also a demand for a revision of the species and genus concept in the Bromeliaceae, taking into account morphological and anatomical as well as molecular data. In the course of such a revision, the taxonomic relevance of commonly used morphological descriptors may have to be re-evaluated.
Descriptions and kty Fascicularia Mez in Mart. Fascicularia Mez in Mart., Fl. Bras. 3,3: 627 (1894);Mez in DC., A. & C., Monogr. Phan. 9: 7 (1896); Harms in Engler & Prantl, Nat. Pflanzenfam, ed. 2, 15a: 132
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(1930); Mez in Engler, Pflanzenr. 4,32: 5 (1934); Smith & Looser, Rev. Univ. 18,8: 1076 (1934);Muiioz, Fl. silvestres Chile: 151 (1966);McClintock, Watsonia 10: 289 (1974-75); Smith & Downs, Fl. Neotrop. 14,3: 17 1 1 (1979); Weber, J. Brom. SOC. 34,6: 265 (1984); Marticorena & Quezada, Gayana 42: 81 (1985);Benoit, Red List Chilean Terrestrial Plants: 1 19 (1 989); Rauh & GroB, Bromelien: 38 1 (1990);Nelson & Zizka, New Plantsman 4,4: 233 (1997). Type: Fusciculuria bicolor (Ruiz & Pav.) Mez in DC., A. & C.
Bromelia auct. non L., p.p.; Ruiz & Pavon, Fl. Peruv. 3: 33 (1802); Molina, Sagg. Chil. ed. 2: 284 (1810), Gay, Hist. bot. Chil. 4: 9, t. 68 (1853); Morren, Belgique Hort. 23: 229, t. 14 (1873);Morren, Belgique Hort. 26: 161-162, pl. l ( r l 1 (1876); Witte, Sempervirens 47: 553-557 (1889); Wittmack, Gartenflora 39: 345 (1890) Billbergu Schult. f. non Thunb., p.p. in Roem. & Schult., Syst. 7,2: 1255 (1830) Hechtia Rivitre non Klotzsch, p.p.; Journ. SOC.Centr. Hort. France: 361 (1871); Morren, Belgique Hort. 26: 161 (1876); Wittmack, Gartenflora 39: 345 (1890) Rhodostach_ysauct. non Phil., p.p.; Bentham in Bentham & Hooker, Gen. Plant. 3,2; 662 (1883); Wittmack in Engler & Prantl, Nat. Pflanzenfam. 2,4: 45 (1 888); Baker, Handb. Brom.: 27 (1889). Terrestrial or epiphytic herbs, stemless or with short, stout stem. LEAVES numerous, densely rosulate, not forming a distinct cup; sheaths 3-7.5(-9) x 1-3(-4.5) cm, broadly ovate to triangdar-ovate, sometimes narrowed at about the middle, coriaceous in upper half, whitish, membraneous and conspicuously nerved toward base; i-appressed-lepidote, glabrescent toward base; margin spinose-serrate toward apex; blades (1 7 -)22-113( - 133) x (0.6 -)0.8-1.8( - 2.2) cm, linear, gradually attenuate toward a pungent apex, rigid, coriaceous, recurving, channelled or plane above, margin spinose-serrate, spines 0.1-0.25 cm long, antrorsely curved, only basally spreading to retrorsely curved; densely lepidote with coarse spreading scales below, _+ densely lepidote basally to glabrate apically above; finely longitudinally striate below; innermost leaves turning scarlet or red when rosette is in flower, with leaf-blades gradually reduced. Outer bracts ovate to oblong, with very short or without blades, as long as or exceeding the flowers; margin conspicuously serrate apically, inconspicuously serrate basally, densely lepidote below in upper third, _+ lepidote to glabrate toward base; approaching in shape and texture the floral bracts of the outer flowers. INFLORESCENCE (3.3-)4-9 x (3-)4-7.5( - 9.5) cm, capitate, terminal, sessile, f globose, simple, many-flowered, dense. Floral bracts (2.2 -)3-5.8( - 7) cm long, shorter than the developed flowers, ovate to oblong (the outer) to spatulate, rarely ligulate (the inner), the claw 0.35-0.5( -0.7) cm wide, f linear, often keeled, the blade 0.5- 1.1 cm wide, roundish to broadly elliptical, apically acute to obtuse, membraneous, margin irregularly serrate apically, entire or inconspicuously serrate basally; densely lepidote in apical third abaxially, glabrate to inconspicuously lepidote adaxially. FLOWERS (2.2 -)3.3-6.5( - 7.5) x 0.6-0.9( - 1.2) cm, bisexual, actinomorphic, sessile, perfect, with conspicuous, cylindrical epigynous tube. Sepals 3, 1-2( -2.4) x 0.35-0.8 cm, free, coriaceous, carinate, the anterior sepal sometimes f ecarinate, ovate to oblong, retuse or praemorse, with a short inconspicuous mucro; densely lepidote-tomentose abaxially, glabrate toward base, inconspicuously to glabrate adaxially. Petals 3, 1.5-2.6 x 0.4-0.8 cm, free, elliptic, obtuse, fleshy, deep to light blue, bearing 2 scale-like appendages at base, these 0.05-0.15 x 0.03-0.1 cm. Stamens 6, included (not exceeding corolla);
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filaments free, broadened toward base (to 0.15 cm wide); anthers linear, elongate, dorsifixed near base. Pollen globose, irregular-monocoplate,c. 40.5 x 27.0 pm (Ehler & Schill, 1973; Erdtman & Praglowski, 1974). Ovary inferior, 3-locular, dorsally compressed, triangular to narrowly bialate, obconical; style elongate, included (not exceeding corolla), with 3 stigmatic branches, these not contorted; placentae central; ovules anatropous, numerous, not or inconspicuously appendaged, f ellipsoid. FRUIT a berry, up to 7.4 cm long (includingpersistent sepals). Seeds 0.22-0.3 x 0.1-0.12 cm, ovoid to fusiform, glabrous, without appendages, dark brownish, surface rugose.
Chromosome number. No information traced. Distribution. One species, endemic to Central and Southern Chile. Comments. The genus is best distinguished from the vegetatively similar genus Ochuguviu by the relative length of style and stamina (Fasciculuriu: included in the flower; Ochuguviu: emerging), shape of sepals (Fasciculariu: retuse or praemorse, with short inconspicuous mucro; Ochuguviu: acute with pungent apex), appendages and colour of petals (Fasciculuriu: appendages present; petals blue to violet; Ochuguviu: appendages absent; petals rose), and pollen characters (Fasciculuriu:irregularly monocolpate; Ochuguviu: monocolpate). The septal nectaries of E bicolor are of special interest: Although the flowers are clearly epigynous as characteristic for Bromelioideae, the morphology of the septal nectaries approaches that of genera from Pitcairnioideae (&/nu, Deuterocohniu, Abromeitiellu). This might point toward Fasciculuriu representing a more primitive genus among the Bromelioideae (Bohme, 1988). Fmciculuriu bicolor (Ruiz & Pav.) Mez in Mart. Bromeliu bicolor Ruiz & Pav., Fl. Peruv. 3: 33 (1802); Molina, Sagg. Chil. ed. 2: 284 (1810); Gay, Hist. bot. Chil. 4: 9, t. 68 (1853); Morren, Belgique Hort. 23: 229, t. 14 (1873). Type: Chile, regibn VIII, ad portum Coronel, leg. Ochsenius s.n. (BR, neo). Billbe+ bicolor (Ruiz & Pav.) Schult. f. in Roem. & Schult., Syst. 7,2: 1255 (1830). Rhodostuchys bicolor (Ruiz & Pav.) Benth. in Benth. & Hook. f., Gen. Plant. 3,2: 662 (1883); Baker, Handb. Brom.: 27 (1889); Wittmack in Engler & Prantl, Nat. Pflanzenfam. 2,4: 45 (1888). Fasciculuriu bicolor (Ruiz & Pav.) Mez in DC., A. & C., Monogr. Phan. 9:9 (1896); Harms in Engler & Prantl, Nat. Pflanzenfam., ed. 2, 15a: 132 (1930); Smith & Looser, Rev. Univ. 18,8: 1076 (1934);Rauh, J. Brom. SOC.35,4: 213, fig. 13 (1984); Marticorena & Quezada, Gayana 42: 81 (1985). Description. As for the genus. Illustrations. Morren (1873: pl. 14); Wright (1906: pl. 8087), Mufioz (1966: pl. 28) (the figures listed here could not be assigned without doubt to one of the subspecies). Bstribution. Chile, region V to region X. No specimen collected by Ruiz & Pavon could be traced, therefore a neotype was selected (Nelson & Zizka, 1997). Discussion. Three combinations in Rhodostuchys (R. bicolor, R. joinvillei, R. pitcuirnigoliu) are ascribed here to Bentham (in Bentham & Hooker, 1883)(contrary to the opinion
REVISION OF THE GENUS FASCZCUIARL4
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of Smith & Downs, 1979), although two of the new combinations do not explicitly appear in the text. Citation of the basionyms with the place of publication (indirectly for B. bicolor) under the genus Rhodostachys, with a short diagnosis and the remark “Species editae 6 v. 7, , .. quarum nobis adsunt 3. . . .” (Bentham & Hooker, 1883: 662), is regarded as sufficient for valid publication. The three combinations published by Mez in Martius (Fl. Bras. 3,3: 627, 1894; F. bicolor, R pitcaimiijilia, R litoralis) are not validly published, for the citation of the basionyms without authors does not fulfill requirements for indirect reference to a previously published description or diagnosis (ICBN, art. 32.5). The differences in geographical distribution (Fig. 2), habitat and leaf morphology (the latter was observed to be constant in cultivated material), led us to assign to the two groups the rank of subspecies.
1. Leaves succulent especially toward base, adaxial (upper) surface flat, margins not recurved, blade (1.O) 1.2- 1.8(- 2.2) cm wide. . . . . . F. bicolor subsp. bicolor 1* Leaves not markedly succulent toward base, adaxial (upper) surface distinctly channelled, margins distinctly recurving, blade (0.6-)0.8-1.1(- 1.4) cm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. bicolor subsp. canaliculata Fascicularia bicolor subsp. bicolor Fascicularia bicolor subsp. bicolol; Nelson & Zizka, New Plantsman 4,4: 238 (1997). Bromeliajoinvillei E. Morren, Belgique Hort. 26: 161-162, pl. 10-1 1 (1876); Witte, Sempervirens 47: 553-557 (1889); Wittmack, Gartenflora 39: 345 (1890). Type: Jardin botanique Liege, 15 XI.1875, Herbier E. Morren (LG, lecto; hic designatus). Rhodostachys joinvillei (E. Morren) Benth. in Benth. & Hook. f., Gen. Plant. 3,2: 662 (1883); Baker, Handb. Brom: 28 (1889). Rhodostachys micrantha Phil., Anal. Univ. Chile 91: 610 (1895-1896). Type: Chile, in praedio Colcura prope oppidum Lota, Martio 1884, Philippi s.n. (‘leg?) (SGO, holo, not seen; FR photo). Fascicularia micrantha (Phil.) Mez in Engl., Pflanzenr, 4,32: 7 (1934);Smith & Downs, Fl. Neotrop. 14,3: 1713 (1979); Rauh, J. Brom. SOC.35,4: 213 (1984);Marticorena & Quezada, Gayana 42: 8 1 (1985); Benoit, Red List Chilean Terrestrial Flora: 1 19, 120 (1989). FasciculariapamJora Mez in DC., A. & C., Monogr. Phan. 9: 12 (1896). Type: Chile, prope Lota, leg. Philippi s.n. (B, probably destroyed). Fmcicularia bicolor (Ruiz & Pav.) Mez in DC., A. & C., Monogr. Phan. 9:9 (1896) and in Engl., Pflanzenr. 4,32: 7 (1934) p.p. (Hartwig 337, Ochsenius s.n.; non Lechler 434, Philippi 204, 1210 (= ssp. canaliculata), non Gay 155 (= OchagaVia). Leaf-sheaths 4-6( - 9) x 2-3( -4,5) cm; leaf-blades succulent especially toward base due to thick accumulation of colourless water-storage cells adaxially, adaxial (upper) surface flat or slightly convex, 22-60( - 80) x (1 .O -) 1.2-1.8( - 2.2) cm. Inflorescence 4-9 x 4-7.5( -9.5) cm; floral bracts (3.2 -) &5.8( - 7) cm long. Flowers 3.7-6.5( - 7.5) cm long.
Illustrations. Morren (1876: pl. 10,ll); Rauh & GroB (1990: Farbbild 92); Nelson & Zizka (1997: 233). htribution. Chile, region V (San Antonio) to region X (northern Chilot). 0-75 m as. 1.
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Habitat. Principally saxicolous near sea-shore. Comments. Only a photograph of the type of Fascicularia micrantha (Philippi s.n., SGO) could be studied. Although in the protologue the plant is described as having comparatively small and narrow leaves and small flowers, the specimen is regarded as belonging to ssp. bicolor on the basis of leaf shape and habit (the protologue includes also the confusing statement, “. . . Las brhcteas son . . . como 10s petalos de un hermoso color rosado”, pointing toward Ochagavia). Distribution also points strongly toward ssp. bicolor. Other specimens studied (partly collected by Philippi) near the type locality (Lota) have been determined as ssp. bicolor. In l?parvz$ora Mez, the case is even less clear. The type specimen (Philippi s.n., B) seems to have been destroyed. According to the protologue, Mez based RparviJora on a type specimen with small flowers and short, comparatively narrow leaves (“. . . 0,25 m. longa, haud ultra 10 mm lata. . .”). From the description, it is not possible to assign the described plant without doubt to a subspecies. The distribution (Lota) clearly points toward ssp. bicolor. Specimens seen. Chile; V region. Valparaiso: San Antonio, Landbeck s.n. (SGO). VII region. Talca: Playa de Constitucion, Parra 63 Rodrieez 156, 2.iv.1975 (CONC); Constitucion, 72’15%’ 35’20’S, R. (= Reiche) s.n., 6.vi.1890 (SGO) (?); Cauquenes: Chanco, Badilla. s.n. v. 1964 (SGO); Curanipe, Goncalez s.n., ix.1966 (SGO). VIII region. Concepcion: Coronel, Ochsenius s.n., 1866 (GOET); Buchenau s.n., 1866 (BR); Lota, Philippi s.n., iii.1884 (SGO);Hualpen, Gunckel 22358, l.i.1941 (CONC); Concepcion, “Ramunchito”, kleiner Badestrand, ,@ka 1414, 22.i. 1991 (FRP); Talcahuano, Peninsula Tumbes, Junge 5477, 8.viii. 1935 (CONC). Arauco: Lebu, R. ( = Reiche) s.n., i. 1902 (SGO). X region. Valdivia: Corral, Gunckel 609, 3.i. 1929 (CONC); 9623, 16.xii.1928 (CONC); Looser s.n., 22.iii. 1933 (NY);Corral, San Carlos, Gunckel 2 131, 6.iii. 1931 and 5692, 16.v.1935 (CONC); Curiiianco, Gunckel449, 2.ii. 1923 (CONC); without precise locality, G q 250 (P).Osorno: La Barra del Rio Bueno, Sparre 4593,2 1.ii. 1948 (S, SGO). Llanquihue: Cochamo, Morrison 17567, 2 1.iii. 1939 (G, MO). ChiloC: Peninsula Lacui, Iter Talcan, Skottsberp Ancud, Pennell 12561, 6-7.ii.02.1925 0; 364b, 12.vii.1908 (S). Chile, without precise locality: Hurtwk 337 (P); Herbarium Hamburgense s.n. (HBG, 2x); Ball s.n. (MO, Foto); Reed s.n. (K, not seen, NY Foto); Sharmann s.n., 1870 (G). Cultivated material: Jard. Bot. Likge, Morren s.n., 15.xi.1875 (LG); Palmengarten FrankfurtIMain, ex BG Edinburgh (696832), Xizka 1790, 1;x. 1996 (FR). Fascicularia bicolor subsp. canaliculata Fascicularia bicolor subsp. canaliculata E.C. Nelson & Zizka, New Plantsman 4,4: 238 (1997). Type: Chile, region X, Valdivia, La Union, Llancacura; leg. Sparre, B. 3546, 20. - 24.xii. 1947 (S, holo). [Pourretiujoinuillei Chantin ex Rivikre, Journ. SOC.Centr. Hort. France: 360 (1871); Morren, Belgique Hort. 26: 161 (1876);Wittmack, Gartenflora 39: 345 (1890)l. Hechtiu joinvillei Rivikre, Journ. SOC.Centr. Hort. France: 36 1 (1 87 1); Morren, Belgique Hort. 26: 161 (1876) (“Hechtia Joinvillei Hort. nonn. Galliae”); Wittmack, Gartenflora 39: 345 (1890).
REVISION OF THE GENUS FASCICULARL.1
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Bromelia albobracteata Steud. ex Lechl., Berber. Am. Austr.: 53 (1857); nom. nud. (based on Lechler 434). Rhodostachys albo-bracteata (Steud. ex Baker) Baker, Handb. Brom.: 28 (1889);Philippi, Anal. Univ. Chile 91: 61 1 (1895-1896). Type: Chile, North Chile, on trees and rocks, 1852; Philippi 204 (G, GOET, WRSL). Rhodostachyspitcaimiijilia var. kirchhofina Wittm., Gartenflora 39: 345, pl. 1325 (1890) (“pitcaimiaefolia”). Type: Plate 1325 in Gartenflora 39 (1890) (lecto, hic designatus), accompanying description @pa345-347). Fmcicularia kz’rchhofina (wittm.) Mez, Repert. Sp. Nov. 16: 2 (19 1 9); Harms in Engler & Prantl, Nat. Pflanzenfam., ed. 2, 15a: 132 (1930); Mez in Engler, Pflanzenr. 4, 32: 6 (1934); Muiioz, F1. silvestres Chile: 151, lam. 28 (1966); Smith & Downs, Fl. Neotrop. 14,3: 1 7 12 (1979);Rauh, J. Brom. SOC.35,4: 2 13,fig. 12 (1 984);Marticorena & Quezada, Gayana 42: 8 1 (1985); Benoit, Red List Chilean Terrestrial Flora: 1 19, 120 (1989). Fascicularia bicolor (Ruiz & Pav.) Mez in DC., A. & C., Monogr. Phan. 9: 9 (1896) and in Engler, Pflanzenr. 4,32: 7 (1934) p.p. (Lechler 434, Philippi 204, 12 10); non Hartwig 337, Ochsenius s.n. (= ssp. bicolor), non Gay 155 (= Ochagavia). Fmcicularia bicolor (Ruiz & Pav.) Mez sensu Lucibell, Bromeliads 4, 20: 140, pl. 141 (1974); sensu Weber, J. Brom. SOC.34,6: 265, fig. 11 (1984). Leaf-sheaths 3-6( -9) x 1-2( - 3) cm; leaf-blades not markedly succulent toward base, adaxial (upper) . surface distinctly channelled, (1 7 -)25-113( - 133)x (0.6-)0.8-1.1(- 1.4) cm. Inflorescence (3.3-)4-5.5 x (3-)4-6.5(-8) cm floral bracts (2.2 -)3-3.8 cm long; flowers (2.2 -)3.3-4.4 cm long.
Illustrations. Wittmack (1890: pl. 1325); Nelson & Zizka ( 1 997: 232). Distribution. Chile, region VIII (Cobquecura) to region X (ChiloC: Piruquina, Trumao). (20-) 150-900 m a s . 1. Habitat. Principally epiphytic in primary, rarely secondary evergreen forest (“Valdivian forest”). Comments. As far as the typification of R. pitcainziijilia var. hrchhofiana is concerned, no herbarium specimen is cited in the protologue. Moreover, no specimen annotated by Wittmack could be traced in Berlin (B). Therefore, the plate accompanying the protologue is regarded as the type. The plate as well as the description make clear that the plant had deeply channelled leaves. Pourretiajoinvilleiand Hechtiajoinvillei were published by Riviirre ( 187 l), accompanied by a very short description, including characteristic features of Fascicularia bicolor ssp. canaliculata (“. . . feuilles canalicultes colorkes, . . . une inflorescence compacte, composte d’un assez grand nombre de fleurs tubuleuses, colortes en beau bleu, . . .”; Riviirre, 1871: 360). Therefore, the names are here included in the synonymy of I.: bicolor ssp. canaliculata in spite of the fact that no type specimen or illustration has been traced. The description given was based on a plant provided by the horticulturist M. Chantin, who, according to Rivitke, proposed the name Pourreha joinvilhi. Nevertheless, in his publication Riviirre did not definitely associate the final epithet with the genus Pourretia, therefore Fjoinvillei Chantin ex Rivi6re was not validly published. Specimens seen. Chile: VIII regi6n. Nuble: Cobquecura, Ledezma 236, 8.x. 1954 (CONC) (cf. canaliculata).
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IX regibn. Cautin: Temuco, Z 72’55’W 38”50’S, Montero 1814, 15.v.1934 (CONC). X region. Valdivia: La Unibn, Fdo. Catamutun, Behn s.n., 27.x.1928 (CONC); reserve forestikre Valdivia, Billkt 3731, 19.i.1985 (BR); Chaihuin, Gunckel 149, 3.i. 1923 (CONC); Coreltut, Hollennayer 547a, 6.iv. 1922 (CONC); Monument0 Natural Alerce Costero, Gurdner G’Knees 410 1, 7.ii. 1988 (E); Panguipulli, Lintre, H o l h a y e r 547b, 25.i. 1927 (B); Hueicolla, 73’35‘W 40” lo’s,. Landrum 4505, 14.iv.1982 (SGO, NY); San Jose de la Mariquina, Montero 5903, xi.1943 (CONC); La Unibn, Llancacura, Spurre 3546, 20-24.xii. 1947 (S, SGO); Calafqutn, Comber 1015, 17.i.1927 (E); without precise locality, Andreus 745, 3.v.1938 (B); LRchler 434, iii.1854, “prope Arique” (BR, G, P, WRSL); Philippi s.n., 1861 (G, SGO). Osorno: Islote Rupanco, C o d y s.n., iii.1978 (SGO); Pichil, 73”03’W 40”40’S, Rudolph s.n., 9.v.1937 (NY); Cord. de La Reina, Spurre 4587, 20.ii.1948 (S). ChiloC: Piruquina, 73’48’W 42”24’S, Junge 31, 17.ix.1931 (B, CONC); Junge 31 & 305 (one sheet), 17.ix.1931 & 17.02.1932 (B, MUC); Junge 31 & 390 (one sheet), l.iv.1932 (MO); Junge 2205, 17.xi.1932 (CONC); Trumao, 73”1 1W 40’2 l’S, Junge 390, 1 .iv. 1932 (MO, MUC); without precise locality, Skottsbq 364a, 22.vii. 1908 (S). Chile, without precise locality (or locality unidentified): Andreus 726, 4.iv. 1938, “San Juan de la Costa, Fundo Yerba buena” (B); Lechler 132 (US); hyboldt 1861 (M);Niemyer 365 (HBG); Philippi 204, v. 1852 (G, GOET, 0, WRSL); 1210, v. 1852 (B); Fdo. San Martin, Woerner s.n., viii. 1952 (HBG). Cultivated material. Hortus Regimontanus (Konigsberg),Mez s.n. (89-7), 1925 (B); Mez s.n. (89-8) (B); Jard. du Luxembourg d Paris, Morren s.n., ix.1879 (LG); Palmengarten Frankfurt/Main,
