genus Ochagavia Phil.

Literature references:

Morren Icons: kew 24-157* Smith & Downs 1979: 1527

*move your mouse pointer over the page numbers to see comment

Comments:

• Revision of the genus Ochagavia (Bromeliaceae, Bromelioideae)
  Abstract
  Zizka, G., Trumpler, K. & Zollner, 0. Revision of the genus Ochagavia (Bromeliaceae, Bromelioideae). - Willdenowia 32. 331-350.2002. -ISSN 0511-9618.
  
  The genus Ochagavia is revised. Four species, 0. andina, 0. carnea, 0. elegans and 0. litoralis, are recognized, all are endemic to Chile. A key to the species, descriptions, illustrations, full synonymies and data on the distribution and ecology are given. The new combinations 0. litoralis and 0. andina are validated, and several names are typified. Relationships of the genus within the subfamily Bromelioideae are discussed.
  
  Introduction
  The Bromeliaceae of Chile are characterized by high endemism, 20 of the 23 species reported for the country being endemic. Only among Tillandsia do we find two species (T. marconae W. Till & Vitek and T. virescens, Ruiz & Pav.) with a wider distribution extending to the arid regions of adjacent Peru and Bolivia, and additionally the widespread T. usneoides (L.) L.
  The genus Ochagavia has a limited distribution in central Chile, ranging from 31°33' to 38°14'S. The attractive plants have been introduced early into cultivation in Europe and appear to grow outdoors as escapes from cultivation in the western parts of Europe with oceanic, mild climate. The morphological similarity with the monospecific genus Fascicularia Mez, especially of sterile plants, has been the reason for some nomenclatural confusion.
  The genus Ochagavia was described by Philippi (1856) originally as monospecific, comprising only 0. elegans Phil. Little later Philippi ( 1858) described the new genus Rhodostachys comprising R. andina Phil. (= 0. andina) and, subsequently, R. litoralis Phil. (= 0. litoralis). Nevertheless, delimitation between Ochagavia and Rhodostachys (and Fascicularia) remained unclear. Finally, Mez ( 1896) realized the close relationship of 0. elegans and Rhodostachys and united the two genera, erroneously giving the later name Rhodostachys priority. Later, Smith & Looser (1934) and Mez (1935) corrected this mistake. For the nomenclature of Fascicularia see Nelson & Zizka ( 1997) and Zizka & al. ( 1999).
  Ochagavia can best be distinguished from Fascicularia by the relative length of the style and stamens, by sepal shape, petal appendages, petal colour and pollen morphology. Although flowering plants of the two genera display striking differences, the relevant characters are difficult to study in herbarium specimens. Sterile specimens are sometimes hard to assign to one of the two genera.
  Our studies are based on herbarium material ( 121 specimens, including duplicates), living collections (7 accessions, including 0. carnea, 0. litoralis and 0. elegans) and observations in the natural habitat.
  
  Morphology, anatomy and cytology
  The representatives of Ochagavia are caulescent, rosulate terrestrials. Although especially in 0. carnea the inner leaves are more or less erect and grouped densely together, the plants do not form tanks that hold water. 0. elegans and 0. litoralis, in particular, have the noteworthy ability to form dense colonies through vigorous offset production. This enables the plants to cover considerable areas, especially on coastal rocks (Fig. 4, 6). The leaves are succulent with spinose-serrulate blades.
  The simple, terminal inflorescence is shortly pedunculate (Fig. 5, 7). The sepals and petals (Fig. 10) are free, the latter not appendaged and rose-coloured. The stamens are exserted (except in 0. elegans), an important character to distinguish the genus from Fascicularia. Another noteworthy character is the well developed epigynous tube, which reaches a length of up to 1.9 cm in 0. elegans.
  Reliable features to separate the Ochagavia species are the indumentum and the size of the leaf blade (Fig. 1). 0. carnea is characterized by long leaves, while 0. elegans has the shortest ones. 0. andina and 0. litoralis are more difficult to separate, but the leaves of the former are longer and narrower than those of the latter. .
  Various authors have used floral characters to distinguish the species of Ochagavia, but our morphological studies, based on herbarium material as well as living plants cultivated in the Palmengarten in Frankfurt am Main, revealed a high variability of these features. Examples are the sepal length, used, e.g., by Smith & Downs (1979), and the petal length; their variability is shown in Fig. 2. Exceptions are the extraordinary length of the epigynous tube and the stamens equalling the petals as most suitable features to characterize 0. elegans. The petals of all Ochagavia species are, to our knowledge, rose-coloured; reports of yellow petals in 0. carnea (e.g. Smith & Downs 1979, as 0. chamissonis ) are erroneous. Also the shape of the bracts, used by Mez ( 1896) to characterize species, has no taxonomic relevance.
  The leaf blade anatomy of Ochagavia litoralis has been studied by Horres & Zizka ( 1995), there erroneously cited as 0. carnea. The considerable water storage tissue comprises 66 % of the leaf (measured as percentage of area of transverse section), making that species one of the most succulent of the Bromeliaceae, comparable to species of the Pitcairnioideae such as Puya roezlii E. Morren (Peru) or Hechtia glabra Brandegee (Mexico ). The water storage tissue is well developed only adaxially, as typical for Bromelioideae and most Pitcairnioideae .
  In Ochagavia carnea a chromosome number of 2n = 50 was counted (Benko-Iseppon & Horres, pers. comm.). The base number x = 25 is predominant in Bromelioideae and in most of the investigated Bromeliaceae (Smith & Till 1998).
  
  Geographical distribution and ecology
  
  Ochaguvia occurs from 31°33' to 38°14'S, being principally restricted to central Chile with a more or less Mediterranean type of climate. Grau ( 1995), following Schmithusen ( 1956), discerns two principal vegetation zones in this area: the "region de bosque esclerofilo" (sclerophyllous forest) in the north and "region del bosque caducifolio templado" (temperate deciduous forest) in the south. In contrast to Fascicularia (occurring from 34° to 42°24'S), Ochagavia does not reach the Valdivian forest zone (Fig. 3), contrary reports are in our opinion erroneous (see a corresponding note by Zizka & al. 1999 regarding a specimen of' Fascicularia pitcairnifolia' = 0. litoralis, leg. Fricke, which is probably referable to F. bicolor) .
  The four Ochagavia species differ with respect to habitat and ecology. 0. elegans from Isla Robinson Crusoe of the Juan Fernandez archipelago typically grows at elevations of 200-600 m on exposed rocks and steep cliffs, forming large stands there. 0. litoralis from continental Chile is also saxicolous but grows close to the sea, at elevations of 10-250 m. Knowledge about the habitat and distribution of 0. andina is scarce; obviously, the species occurs further inland, extending from 700-2500 m elevation. The remarkable, large plants of 0. carnea are found from Valparaiso to Malleco between (60- )200 and 900 m elevation. Specimens have been collected from moist understorey of woodlands and in ravines, especially near river banks. Apparently, the latter species is the one in the genus that prefers the most humid habitats.
  While Chilean Greigia species (G. pearcei Mez and G. berteroi Skottsb.) appear to have become very rare, and the latter being possibly in danger of extinction (Will & Zizka 1999, Danton 2002), the situation in Ochagavia and Fascicularia is different. In contrast to G. berteroi, the only other bromeliad in the Juan Fernandez archipelago, 0. elegans is still quite abundant on rocky cliffs of Isla Robinson Crusoe. Nevertheless, because of the various threats for this unique island flora, Hoffmann & Flores ( 1989) regard this species as "vulnerable". 0. litoralis is in continental Chile comparatively abundant too. No information about population size and possible endangerment is at hand for the two other species that occur further inland. The fact that the known specimens of 0. andina were collected many decades ago indicates that this species may have become very rare.
  
  Systematic relationships and phytogeography
  The morphological similarity between Ochagavia and Fascicularia points toward a close relationship, which is supported by investigations using molecular markers (Horres & al. 2000, Horres & al. in prep.), While RAPD studies revealed also a close relationship with Greigia sphacelata (Ruiz & Pav .) Regel (Zizka & al, 1999), this could not be supported by sequence data from trnL intron (Horres & al. 2000, Horres & al. in prep.). The latter study included 16 genera of Bromelioideae and revealed interesting outcomes: despite their close geographical association, Greigia and Fascicularia/Ochagavia occupy quite different positions in the phylogenies obtained. Whereas Ochagavia and Fascicularia form a clade together with the monospecific Androlepis skinneri Brongn. ex Houllet, which grows epiphytically in Central American and Peruvian forests, Greigia groups together with Wittrockia and Lymania, both being endemic to southeastern Brazil.
  The systematic affinities of Ochagavia, Fascicularia and Greigia are still not clear, This holds true for the entire Bromelioideae. Due to the low genetic variability observed in the markers studied up to now, further evidence is needed to produce more reliable phylogenetic reconstructions. Nevertheless, monophyly of this subfamily has been confirmed by all molecular studies at hand.
  The southern border of the distribution area of the Bromeliaceae west of the Andes is situated at 42°24'S (the record of Fascicularia bicolor (Ruiz & Pav ,) Mez from 45° 17'S could not be confirmed by us, see also Zizka & al. 1999) and east of the Andes at 44°46'S, On either side of the Andes, species and genera of different subfamilies form the southern limit. In the east, the southernmost bromeliads are representatives of the Tillandsioideae, these are Tillandsia pedicellata (Mez) A. Cast., T. retorta Griseb. ex Baker and T. andicola Gillies ex Baker, the first being reported as far south as 44°46'S in the province Chubut, southern Argentina. All three species have a comparatively large distribution area in semidesert and steppe climates, extending to northwestern Argentina or, in the case of T. pedicellata, even to western Bolivia (Till 1984, Till, pers. comm,). West of the Andes, in the area of the Valdivian rain forest, the southermost bromeliads are represented by Greigia, Fascicularia and, further north, Ochagavia, all belonging to the Bromelioideae. The species occurring there are endemics, in the case of Fascicularia and Ochagavia even restricted to this comparatively small area,
  
  
  Cultivation
  Plants of Ochagavia are capable of being cultivated outdoors in (north)western Europe. According to Clement & Foster (1994) and Stace (1997: 922), 0. carnea has become naturalized on Tresco, Isles of Scilly ("Tresco Rhodostachys", "... well established where planted on dunes in Tresco, Scillies"). 0. carnea and 0. litoralis, recently also 0. elegans (Wilkin 1996), are cultivated in botanical gardens, usually together with succulents. As observed in the Palmengarten, Frankfurt am Main, 0. carnea plants very rarely flower, less often than the Fascicularia plants grown under similar conditions. Nevertheless, Ochagavia species are attractive plants and can be recommended for cultivation, especially the smaller ones, 0. litoralis and 0. elegans. The latter species, up to now rarely in cultivation, might even prove to be hardy outside in western Europe, but, as reported by Wilkin ( 1996), is not frost hardy.
  
  
  Chromosome number. - 2n = 50 (Benko-Iseppon & Horres, pers. comm.).
  
  Distribution. - Three species endemic to continental central Chile, one endemic to Isla Robinson Crusoe, Juan Fernandez archipelago (Fig. 3).
  
  Remarks. - The genus is best distinguished from the vegetatively similar, monospecific genus Fascicularia by the following features:
  - relative length of style and stamina (Ochagavia: exserted, Fascicularia: included in the flower);
  - shape of sepals ( Ochagavia: acute with pungent apex, Fascicularia: retuse or
  premorse, with short inconspicuous mucro); .,
  - colour and appendages of petals ( Ochagavia: petals. rose, appendages absent,
  Fascicularia: petals blue to violet, appendages present);
  - pollen characters (Ochagavia: monocolpate, Fascicularia: irregularly
  monocolpate) (see also Zizka & al. 1999).
  Morphological and molecular studies (Horres &al. 2000, Horres & al, in prep.) revealed a very closely relationship of the two genera.
  
  
  Further references. - Ochagavia: Philippi in Bot. Z. 14: 647. 1856; F. Philippi in Anales Univ. Chile 59: 278. 1881; Bentham in Bentham & Hooker, Gen. PI. 3,2: 661. 1883; Wittmack in Engler & Prantl, Nat. Pflanzenfam. 2(4): 45.1887-88; Harms in Engler & Prantl, Nat. Pflanzenfam., ed. 2, 15a. 159.1930; Mez in Engler, Pflanzenr. 100: 203.1935; Munoz, Espec. Pl. Descr. Philippi: 35. 1960; Smith & Downs in Fl. Neotrop. Monogr. 14,3: 1527. 1979; Rauh & Gross, Bromelien: 398. 1990; Grant & Zijlstra in Selbyana 19: 105. 1998; Smith & Till in Kubitzki, Fam. Gen. Vasc. PI. 4: 93. 1998. - Rhodostachys: Philippi in Anales Univ. Chile 91: 607. 1895; Baker, Handb. Bromel.: 27. 1889; Mez in C. Candolle., Monogr. Phan. 9: 335. 1896; Reiche, Grundz. Pfl.-Verbr. Chile: 67. 1907; Harms in Engler & Prantl, Nat. Pflanzenfam., ed. 2, 15a: 132, 159. 1930; Munoz, Espec. Pl. Descr. Philippi: 35. 1960; Grant & Zijlstra in Selbyana 19: 109.1998. - Ruckia: Regel in Gartenfl. 17: 65, t. 571.1868; Grant & Zijlstra in Selbyana 19: 109. 1998. - Placseptalia: Grant & Zijlstra in Selbyana 19: 106. 1998.
  
  Key to the species of Ochagavia
  
  1. Epigynous tube more than 10 mm long, stamens not or barely exceeding the petals
  (style distinctly exceeding); endemic to Isla Robinson Crusoe . . . . 1. elegans
  1a Epigynous tube up to 5( - 7) mm long, stamens (as well as style) exceeding the petals;
  endemic to continental Chile . . . . . . . . . . . . . 2
  
  2. Leaf blades 15-45cm long, densely whitish lepidote below . . . . . . 3
  2a Leaf blades (40-)50-80(-120) cm long, sparsely lepidote to glabrescent below.
  4. carnea
  
  3. Leaf blade 15-40 x l-3cm . . . . . . . . . . 2 litoralis
  3a. Leafblade30-45 x 0.9-1.7cm . . . . . . . . . . 3. andina
  
  
  Nomina dubia Ochagavia leiboldiana Rhodostachys leiboldiana Mez in Candolle, Monogr. Phan. 9: 338. 1896 ["leiboldianus"]. Holotype: Chile, without locality ("in Exp. Donau"), Leibold 2960 (W, destroyed) = Ochagavia leiboldiana (Mez) L. B. Sm. & Looser in Rev. Univ. (Santiago) 18(8): 1078. 1934.
  Remarks. - The type specimen deposited in the collections of the Natural History Museum in Vienna (W) was destroyed in a fire shortly after World War II (Till 1994). No collecting data (coastal or Andean habitat) are given in the protologue, thus it remains doubtful from the description, whether Rhodostachys leiboldiana must be regarded conspecific with Ochagavia litoralis or O. andina. For this reason, we did not select a neotype. The information given about leaf size (length: ± 0.4 m, width: up to 8 mm) and indumentum of the abaxial leaf blade surface clearly points toward one of the above mentioned Ochagavia species.
  Further references. -Mez in Engler, Pflanzenr. 100: 204. 1935.
  
  Detail from S&D
  25. Ochagavia R. Philippi, An. Univ. Chile 13: 168. May 1856; Bot. Zeitung 14: 647. Sep 1856. Fig 489.
  Rhodostachys R. Philippi, Linnaea 29: 57. 1857-58.
  Ruckia Regel, Ind. Sem. Hort. Petrop. "1867": 28. 1868 (?); Gartenflora.
  Placseptalia Espinosa, Bol. Mus. Nac. Hist. Nat. Chile 23: 5. 1947.
  Suffruticose very stiff herbs, often more of less erect-caulescent, amply proliferating. Leaves very numerous, not tank forming; blades linear, spinose-serrate. Scape none or very short. Inflorescence sunk in the center of the leaves at the apex of the stem, simple, short, globose or capitate-subracemose, many-flowered. Bracts conspicuous, narrow; flowers perfect. Sepals free, overlapping only at the extreme base, symmetric or subsymmetric, mucronulate; petals free, symmetric, rose or yellow, claw short, blade narrowly elliptic or suboblong, rounded, naked; stamens exserted or rarely only equaling the petals; anthers subelliptic, dorsifixed 1/3-1/2 above the base; pollen eporate, with a single longitudinal fold; ovary inferior, strongly compressed or angled, glabrous; epigynous tube very conspicuous; placentae extending the whole length of the locule; ovules rather numerous, unappendaged; fruit distinctly enlarged from the ovary, crowned by the persistent sepals; seeds large, globose.
  
  Type. Ochagavia elegans R. Philippi, An. Univ. Chile 13: 168. May 1856; Bot. Zeitung 14: 647. Sep 1856.
  
  Key to the Species of Ochagavia
  1. Epigynous tube 10 mm long; leaves distributed along the elongate stem, 10-25 cm
  long; stamens about about equaling the petals; sepals to 18 mm long. elegans
  1. Epigynous tube ca. 5 mm long at most; leaves rosulate, 20-70 cm long; stamens
  conspicuously exserted at anthesis; sepals 12-27 mm long.
  2. Leaves glabrous or subglabrous on both sides; petals yellow; sepals to 19 mm long.
  chamissonis.
  2. Leaves densely white-lepidote beneath; petals rose; sepals 12-27 mm long. carnea