Botanical Journal of the Linnean Society, 2008, 156, 93–110. With 4 figures Morphological phylogenetics of Puya subgenus Puya (Bromeliaceae) CLAUDIA T. HORNUNG-LEONI and VICTORIA SOSA* Departamento de Biología Evolutiva, Instituto de Ecología, A.C. Apartado Postal 63, 91000 Xalapa, Veracruz, Mexico Received 1 November 2006; accepted for publication 3 September 2007 Puya, a large genus mostly from South America, has been taxonomically divided into two subgenera: Puyopsis and Puya. The latter includes only eight species distributed mainly in Chile, extending to Argentina, Bolivia, and Peru. The species of subgenus Puya are recognized by the presence of a sterile apex of the inflorescence branches, whereas those of subgenus Puyopsis have fertile flowers all along the branches. The objectives of this article were to determine whether this diagnostic character was synapomorphic for subgenus Puya, and to explore the relationships between its species. Parsimony analyses were performed for 43 taxa and 93 morphological characters, 87 of which were discrete and six continuous. In the analysis that included all characters, a single most parsimonious tree was found that supported subgenus Puya by two synapomorphic character states, including the diagnostic character of a sterile inflorescence branch apex and a blooming pattern in which flowers open gradually from base to apex. The trees were better supported when the continuous characters were included. Further studies are suggested to resolve the infrageneric classification of Puya and the relationships of the species belonging to subgenus Puya. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110. ADDITIONAL KEYWORDS: continuous characters – morphology – Puya subgenus Puyopsis. INTRODUCTION Puya Molina is a genus of Bromeliaceae that is almost exclusive to South America – only two taxa are found in Costa Rica – and includes approximately 199 species (Smith & Downs, 1974; Luther, 2002). The description of Puya is based on diagnostic attributes, such as petals spiralled and persistent after anthesis (Smith & Downs, 1974). The genus was previously divided into a number of subgenera, including Chagualia, Pitcairniopsis, and Pourretia (Mez, 1896; Smith & Looser, 1935). However, only two subgenera are currently recognized: Puya and Puyopsis (Smith, 1970). Subgenus Puyopsis (Baker) L.B. Sm. has fertile flowers along the branches of inflorescences, whereas subgenus Puya lacks fertile flowers at the apex of the branches of inflorescences (Smith, 1970; Smith & Downs, 1974). *Corresponding author. E-mail: victoria.sosa@inecol.edu.mx The majority of the species belong to subgenus Puyopsis, and only nine have been included in subgenus Puya. Smith & Downs (1974) initially included seven species in subgenus Puya: Puya chilensis Molina (the type species of Puya), Puya boliviensis Baker, Puya castellanosii L.B.Sm., Puya alpestris (Poepp.) Gay, Puya berteroniana Mez., Puya weddelliana Mez, and Puya raimondii Harms. Subsequently, Puya quillotana W. Weber (1984) and Puya gilmartiniae G.S. Varadarajan & A.R. Flores (1990) were described. However, the only differentiating character between P. quillotana and P. chilensis was the pubescence of the leaves recorded from a single incomplete specimen. Therefore, here it is considered as a synonym of P. chilensis, and only eight species are included in subgenus Puya. The morphological variation in Puya is remarkable, mainly in the size of plants and floral parts, the branching patterns of the inflorescence, and the arrangement of the flowers in the inflorescence. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 93 94 C. T. HORNUNG-LEONI and V. SOSA Some of these variable characters are found amongst species of subgenus Puya. For example, the largest bromeliad in the world, P. raimondii, reaches approximately 12 m in height, in comparison with P. alpestris that only grows to approximately 1.5 m. Plants of P. boliviensis lack a stem, whereas the plants of the rest of the species of subgenus Puya have a welldeveloped stem, which may be simple, as in P. raimondii, or branched, as in P. chilensis. The position of Puya has posed a problem in the phylogenetic analyses of Bromeliaceae. It has been included in Pitcairnioideae, related to Pitcairnia by morphological characters and chloroplast DNA restriction site variation (Varadarajan & Gilmartin, 1988; Ranker et al., 1990). However, current molecular approaches that have considered one or a few species of Puya have indicated that the Bromelioideae is the sister group of the genus (Terry, Brown & Olmstead, 1997; Crayn et al., 2000, Crayn, Winter & Smith, 2004; Givnish et al., 2004; Barfuss et al., 2005). The traditional classification of the Bromeliaceae has recognized three subfamilies: Pitcairnioideae, Bromelioideae, and Tillandsioideae (Smith & Downs, 1974). Recent phylogenetic studies have not agreed on the composition or position of these groups; contradictory results are mainly related to which groups emerged first (Clark & Clegg, 1990; Ranker et al., 1990; Terry et al., 1997; Crayn et al., 2000; Horres et al., 2000; Givnish et al., 2004; Barfuss et al., 2005). The monophyly of the Pitcairnioideae has been questioned, and therefore both new subfamilies and tribes could be defined in the future (Ranker et al., 1990; Terry et al., 1997; Givnish et al., 2004). By the way of an example, a new tribe – Puyeae – has been suggested (Terry et al., 1997; Benzing, 2000). Phylogenetic analyses of Puya are still lacking. The only study that evaluated morphological data included only nine species (Varadarajan & Gilmartin, 1988), not sufficient to understand this varied and complex group. Current molecular phylogenetic approaches in the Bromeliaceae have included only one or a few species of Puya (Terry et al., 1997; Crayn et al., 2000, 2004; Givnish et al., 2004; Barfuss et al., 2005). The aims of this study were to carry out phylogenetic analyses based on morphological characters to determine the characters that support subgenus Puya, and to investigate the relationships between the species of this subgenus. Molecular approaches in Bromeliaceae have found very little variation in a number of chloroplast DNA regions, such as trnK, rps16, trnL, trnL-trnF atpB-rbcL, rbcL, and matK (Terry et al., 1997; Crayn et al., 2000, 2004; Givnish et al., 2004; Barfuss et al., 2005), and nuclear regions, such as internal transcribed spacer (ITS), are only just beginning to be explored (Horres et al., 2000). Therefore, this study provides an initial estimate of the phylogenetic relationships of the species of subgenus Puya until variable molecular characters are found. In addition to discrete morphological attributes, continuous characters were considered in the analyses. There is controversy over the use of these characters for cladistic analysis. Arguments against the use of these characters are threefold: (1) those related to the concept of homology; (2) those against the way in which continuous characters are broken down; and (3) those against the concept that characters are classes and not individuals (Archie, 1985; Pimentel & Riggins, 1987; Cranton & Humphries, 1988; Chappill, 1989; Stevens, 1991; Thiele, 1993; Rae, 1998; Kluge, 2003; Grant & Kluge, 2004). In this study, six continuous characters were coded according to their range of measurement, utilizing the computer program TNT (Goloboff, Farris & Nixon, 2003). TNT deals with continuous characters as such, avoiding the use of ad hoc methods that have been proposed to discretize continuous distribution in phylogenetic analysis (gap-coding, Thiele’s method, etc.) (Goloboff, Mattoni & Quinteros, 2006). The inclusion of continuous characters will allow for an understanding of the evolution of the size of several floral and vegetative elements in subgenus Puya. MATERIAL AND METHODS TAXON SAMPLING In addition to the eight species of subgenus Puya, 20 taxa of subgenus Puyopsis were included on the basis of the following criteria: (1) species representative of the entire range of distribution; and (2) species representative of different morphological patterns, well represented in herbaria. For the outgroups, 15 taxa representative of the three traditional subfamilies were selected: Pitcairnioideae, Bromelioideae, and Tillandsioideae (Baker, 1889; Smith & Downs, 1974). These taxa correspond to the different clades identified by molecular phylogenetic studies (Clark & Clegg, 1990; Ranker et al., 1990; Terry et al., 1997; Crayn et al., 2000; Horres et al., 2000; Givnish et al., 2004; Barfuss et al., 2005). Forty-three terminal taxa were included (see Appendix 1). From these, Brocchinia was used to root the tree, as it is the most distantly related genus. MORPHOLOGICAL DATASET The data matrix (Appendix 2) consists of 93 characters: 87 discrete and six continuous characters. Vegetative as well as floral and micromorphological characters were included. Some of these characters are shown in Figures 1 and 2. Micromorphological © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 PHYLOGENY OF PUYA SUBGENUS PUYA 95 Figure 1. Floral and epidermal characters: A, stamens adnate to petals; B, stellate hairs; C, trichomes with central disc cells and wings; D, trichomes without a central disc but with wings. characters were observed with either an optical microscope or a JEOL JSM-5600 LV scanning electron microscope. Four hundred and sixty-six herbarium specimens were examined, some of which were collected for this project. They are listed in Appendix 1. The characters and their states are given in Table 1. For characters 25 and 26, the percentage of hairs on both abaxial and adaxial surfaces was estimated by dividing the microscope field into four. Continuous characters (Table 2) were divided into character states with TNT. The program considers measurements as a range (minimum/maximum) of the character for each species with values from 0 to 65, using up to three decimals. The characters were optimized using Farris’ method for additive characters (Farris, 1970). During the optimization of a given node, if the ranges of the descendant nodes overlapped, the method counted no steps; if the ranges did not overlap, it counted the minimum distance from one range to the other (i.e. the numeric difference between the two closest values of the two descendant ranges) (Goloboff et al., 2006). Measurements were standardized by a log(x + 1) transformation, because of differences in scale (Table 2). There were up to 32 states for some of these characters, as implemented in TNT (Goloboff et al., 2003). PHYLOGENETIC (Goloboff et al., 2003). Separate analyses were performed: the first dataset included only the discrete characters, and the second included the discrete and continuous characters. Parsimony analyses were performed with 1000 starting trees with tree bisection– reconnection (TBR), saving 50 trees per replication. Support was estimated by jackknife by resampling 1000 times with the TBR set with a removal probability of 30%. Bremer support (Bremer, 1994) was calculated only for the combined dataset as implemented in TNT (Goloboff et al., 2003). RESULTS CLADISTIC ANALYSES The analysis with discrete characters retrieved nine most parsimonious trees (MPTs) [L = 505 steps; consistency index (CI), 0.250; retention index (RI), 0.522]. The strict consensus is shown in Figure 3. In the analysis with discrete and continuous characters, a single MPT was retrieved (Fig. 4) (L = 530 steps; CI, 0.250; RI, 0.515). In the data matrix with discrete characters, the 87 characters were parsimony informative. In the data matrix with continuous + discrete characters, 93 characters were parsimony informative. Autapomorphic characters were scored but not taken into account in the analyses. ANALYSIS The data matrix was constructed using WINCLADA (Nixon, 2002). Cladistic analyses under parsimony criteria were performed using the program TNT PHYLOGENETIC RELATIONSHIPS In the discrete character analysis, only a clade that included two Pitcairnia species was well supported © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 96 C. T. HORNUNG-LEONI and V. SOSA © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 PHYLOGENY OF PUYA SUBGENUS PUYA 97 Figure 2. Vegetative and floral characters. Rosette type: 1A, tank; 1B, cistern; 1C, tufted. Stem: 1A, acaulescent; 2A, caulescent. Stem type: 2A, erect; 2B, prostrate. Stem branching: 2A, simple; 2B, branched. Inflorescence/rosette relative length: 3A, same; 3B, once the length; 3C, twice the length; 3D, smaller. Leaf margin: 4A, entire; 4B, serrate. Thorn orientation: 5A, antrorse; 5B, retrorse; 5C, antrorse and retrorse. Peduncle: 3D, included; 3B, emerging; 3A, equalling. Peduncle covered with bracts: 3B, totally; 3C, partially. Inflorescence branching: 6A, simple; 6B, branched. Inflorescence strobiliform: 6B, absent; 3B, present. Flower density: 6A, lax; 7A, dense. Inflorescence branch arrangement: 7B, polystichous; 6A, distichous. Sterile apex of inflorescence branch: 6B, absent; 7B, present. Petal shape: 8A, oblong-elliptic; 8B, obovate. Petals spiralled and twisted: 9A, present. Petaloid appendices at base of petals: 8B, absent; 8C, present. Fruit type: 10A, capsule; 10B, berry. Fruit dehiscence: 11A, septicidal; 11B, loculicidal and septicidal. Petals persistent at apex of fruit: 11C, present. Seed: 12A, naked; 12B, plumose; 12C winged; 12D appendiculate. 䉳 (jackknife 93%; Bremer support value, > 2) (Fig. 3). The species of Puya were grouped in a clade (Bremer support value, > 1). The species in subgenus Puya were grouped in a subclade without support within a larger clade together with Puya goudotiana and Puya retrorsa (Bremer support value, > 1). In the discrete + continuous character analysis, more clades received support (Fig. 4). The Puya clade received a jackknife value of 51% and a Bremer support value of > 3. The single MPT retrieved by this analysis shows species of Puya in two clades without support. The first is the Puya laxa clade and the second has two subclades: the subgenus Puya subclade and the Puya aristeguietae subclade. The subgenus Puya clade received a jackknife value of 52% and a Bremer support value of > 2 (Fig. 4). SYNAPOMORPHIC CHARACTERS In the discrete character analysis, the genus Puya was supported by six synapomorphic character states, two of which were unambiguous: petals spiralled and twisted together after anthesis (77:1; CI, 100) and petals spiralled persistent at the apex of fruit (91:1; CI, 100) (Fig. 3). Subgenus Puya was supported by two character states: presence of a sterile inflorescence branch apex (44:1; CI, 100) and flowers opening sequentially from inflorescence branch base to apex (47:0, CI, 100). In the analysis that included the discrete + continuous characters, the genus Puya and the subgenus Puya were supported by the same synapomorphic states as in the discrete character analysis (Fig. 4). DISCUSSION In this study, the species classified in subgenus Puya were recovered in a clade supported by two synapomorphic character states. It was confirmed that the diagnostic character for the subgenus (presence of a sterile inflorescence branch apex) is a synapomorphy that supports this clade. The other character state that was synapomorphic was the blooming pattern, in which the flowers open gradually from base to apex. Johow (1898) suggested that a sterile apex on the branches of the inflorescence is an adaptation that provides support for perching birds. We have observed in the field that, in P. raimondii, both hummingbirds and passerine birds visit the flowers, and so the sterile apex of branches in this species is not specifically for perching birds. However, more information on the pollination system of the species in subgenus Puya will clarify the importance of the sterile apex in relation to pollinators. Our results imply that subgenus Puyopsis is paraphyletic. The studied species of this subgenus are grouped into two clades, one of which is the sister group of the species of subgenus Puya. Several other subgenera have been proposed in the past, including Chagualia Smith & Looser, Pourretia Mez, and Pitcairniopsis Mez (Smith & Looser, 1935; Smith, 1970), and, most probably, some will have to be resurrected. Thus, further analyses with more taxa and a larger number of characters are needed to recover trees that are better resolved, and to determine the infrageneric classification of Puya and the relationships of its species. Such analyses will also clarify the status of the group formed by northern South American species represented by the P. aristeguietae clade that is closely related to subgenus Puya. The topology of the cladograms was better supported when continuous characters were included. None was synapomorphic. This coincides with the results of previous analyses in which these characters were taken into account (for example, Lehtonen, 2006). We did not break up the range of measurements of the characters, as normally carried out (see Garcia-Cruz & Sosa, 2006). Instead, we used TNT, which analyses continuous characters as such. Our results indicate that continuous characters contain phylogenetic information, and justifies that they are homologous. When the continuous characters are visualized in the single MPT (Fig. 4) retrieved by the discrete + continuous character data matrix, the general pattern of species of subgenus Puya is that of an increase in plant, inflorescence, sepal, and petal size. The species of subgenus Puya are in a grade, with the © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 98 C. T. HORNUNG-LEONI and V. SOSA Table 1. Characters and character states for the cladistic analysis of subgenus Puya 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. Plant length Leaf length Inflorescence length Floral bract length Sepal length Petal length Rosette type: 0, tank; 1, cistern (tubular cylindrical); 2, tufted (graminoid). In the tank type, the leaves are at an angle of 45° and water is retained; in the cylindrical or tubular type, the leaves overlap and they are erect and form a container; the tufted type is similar to the habit of grasses Stem: 0, acaulescent; 1, caulescent Stem type: 0, erect; 1, prostrate Stem branching: 0, simple; 1, branched Reproduction: 0, monocarpic; 1, polycarpic Inflorescence/rosette relative length: 0, same; 1, inflorescence once the length of rosette; 2, inflorescence twice the length of rosette; 3, inflorescence smaller than rosette Leaf position in the rosette during blooming: 0, erect; 1, recurved-reflexed; 2, ascendent Leaf margin: 0, parallel; 1, convergent Leaf blade apex: 0, acute or attenuate; 1, rounded; 2, mucronate or acuminate Leaf capacity of leaves to retain water: 0, absent; 1, present Leaf margin: 0, entire; 1, serrate Leaf margin: thorn orientation: 0, one way, antrorse; 1, one way, retrorse; 2, with both orientations, antrorse and retrorse Leaf margin: thorn colour: 0, black-brownish; 1, brown-reddish or greenish; 2, yellow to red Leaf nervation: 0, without a prominent median nerve; 1, with a prominent median nerve Leaf petiole: 0, absent; 1, present Leaf pubescence of adaxial surface: 0, glabrous; 1, with scales; 2, with hairs (pubescent) Leaf pubescence of abaxial surface: 0, glabrous; 1, with scales; 2, with hairs (pubescent) Leaf adaxial and abaxial surfaces: trichomes: 0, with central disc cells and wings; 1, without a central disc but with wings; 2, irregular cells with neither central disc nor wings Leaf adaxial surface: percentage of pubescence: 0, 25%; 1, 50–75%; 2, 100% Leaf abaxial surface: percentage of pubescence: 0, 25%; 1, 50–75%; 2, 100% Inflorescence: peduncle: 0, included in rosette; 1, emerging rosette; 2, equalling rosette Inflorescence: 0, erect; 1, pendule or nutant Inflorescence: peduncle diameter: 0, less than 2 cm; 1, 2–10 cm; 2, more than 10 cm Inflorescence reddish coloration: 0, absent; 1, present Inflorescence: peduncle covered with bracts: 0, totally; 1, partially Inflorescence: peduncle bracts shape: 0, oblong-elliptic; 1, ovate Inflorescence: bract margin: 0, entire; 1, serrate or serrulate Inflorescence: bract pubescence: 0, glabrous; 1, with scales; 2, with hairs Inflorescence: branching: 0, simple; 1, branched Inflorescence position: 0, apical; 1, lateral Inflorescence: consistency of the rachis of inflorescence: 0, nonfleshy; 1, fleshy Inflorescence: strobiliform: 0, absent; 1, present Inflorescence: flower disposition type: 0, spikes; 1, racemes Inflorescence: flower density in the entire inflorescence: 0, lax; 1, dense Inflorescence: branch arrangement: 0, polystichous; 1, distichous Inflorescence: branch shape: 0, globose; 1, oblong-elliptic Inflorescence: branch position: 0, alternate; 1, verticillate Inflorescence: sterile apex of branches: 0, absent; 1, present Inflorescence: density of flowers along branches: 0, lax; 1, dense Inflorescence: shape: 0, triangular; 1, oblong Inflorescence: blooming pattern: 0, flowers opening sequentially from inflorescence branch base to apex; 1, flowers opening in any position Inflorescence: bract/branch length: 0, bracts smaller or equalling branches; 1, bracts larger Inflorescence: bract length: 0, larger than the axil of an inflorescence’s branch; 1, smaller than or equal to the axil of an inflorescence’s branch © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 PHYLOGENY OF PUYA SUBGENUS PUYA 99 Table 1. Continued 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 88. 89. 90. 91. 92. 93. Inflorescence: bract shape: 0, oblong-elliptic; 1, ovate Inflorescence: apical zone of the inflorescence bracts apex: 0, acute-attenuate; 1, rounded; 2, mucronate-apiculate Inflorescence: margin of bract: 0, entire; 1, serrate or serrulate Inflorescence: blade of bracts: 0, absent; 1, present Inflorescence rachis pubescent: 0, absent; 1, present Inflorescence: ferrugineous pubescence: 0, absent; 1, present Inflorescence: bract apex: 0, acute-attenuate; 2, rounded or obtuse; 2, mucronate or apiculate Floral bract: margin: 0, entire; 1, serrate or serrulate Floral bract: carinate: 0, absent; 1, present Floral bract shape: 0, oblong-elliptic; 1, ovate Floral bract colour (fresh material): 0, greenish; 1, brownish; 2, red–orange Floral sterile bracts among flowers: 0, absent; 1, present Flower disposition: 0, distichous; 1, polystichous Flower phyllotaxis: 0, opposite; 1, alternate; 2, verticillate Flower: pedicel: 0, absent; 1, short (< 2 mm); 2, medium (2–6 cm); 3, large (> 6 cm) Flower: length with regard to bracts: 0, larger; 1, smaller; 2, equal size Flower: bract pubescence: 0, absent; 1, present Flower: sepal length with regard to bracts: 0, larger; 1, smaller; 2, equal size Flower: sepal symmetry: 0, symmetrical; 1, strongly asymmetrical Flower: sepals carinate: 0, absent; 1, present Flower: sepal union: 0, free; 1, sepals connate at least at half; 2, connate only at the base Flower: sepal apex: 0, acute-attenuate; 1, rounded-obtuse; 2, mucronate-acuminate Flower: sepal length with regard to petal: 0, 1/3 smaller; 1, 1/2 smaller or less; 2, equal size Flower: sepal colour: 0, yellowish; 1, reddish or coloured; 2, greenish-white–cream; 3, brownish Flower: sepal pubescence: 0, absent; 1, present Flower: petal colour: 0, white–cream; 1, blue–violet; 2, dark green–blue; 3, red–rose; 4, yellow-greenish Flower: petal shape: 0, oblong-elliptic; 1, obovate Flower: petals spiralled and twisted together after anthesis: 0, absent; 1, present Flower: petal apex: 0, acute-attenuate; 1, rounded-obtuse; 2, mucronate-acuminate Flower: petal symmetry: 0, asymmetric; 1, symmetric Flower: petaloid appendices at base of petals: 0, absent; 1, present Flower: relative size of stamen with respect to the flowers: 0, inserted; 1, exerted Flower: stamens adnate to petals: 0, absent; 1, present Flower: stamens in a column: 0, absent; 1, present Flower: anthers: 0, basifixed; 1, dorsifixed Flower: stamen length with regard to style: 0, larger or equal size; 1, smaller Flower: anther shape: 0, sagittate; 1, oblong-elliptic Flower: ovary: 0, superior; 1, semi-inferior; 2, inferior Flower: style size with regard to ovary: 0, larger; 1, smaller or equal Fruit: type: 0, capsule; 1, berry; 2, drupe; 3, multiple fruit Fruit: dehiscence of capsules: 0, septicidal; 1, loculicidal; 2, both Fruit: petals persistent at apex of fruit: 0, absent; 1, present Seed: 0, naked; 1, plumose; 2, winged; 3, appendiculate Roots: wide radicular system: 0, absent; 1, present exception of a small terminal clade formed by P. castellanosii and P. raimondii. Plants of the former are medium sized, but the latter is the largest bromeliad in the world, with a very large inflorescence (Foster, 1950; Varadarajan & Gilmartin, 1988). In addition, P. raimondii’s flowers have larger petals in comparison with those of the other species of the subgenus, but the inflorescence is gigantic, and thus holds several thousands of flowers. Of all the species in Bromeliaceae, P. raimondii produces the largest number of flowers per inflorescence (Foster, 1950; Varadarajan & Gilmartin, 1988). Puya raimondii does not show clonal growth; it only reproduces by seeds (Hornung-Leoni & Sosa, 2004). Therefore, it is suggested that the reproductive strategy of this species is to produce more flowers to increase seed set. The results of a previous study have shown that there is an allometric pattern in Puya in which plant size is © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 100 Table 2. Continuous characters and their logarithmic transformation 1 2 3 4 5 © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 Species Plant length (cm) Log (height + 1) Leaf length (cm) Log (leaf + 1) Infl. length (cm) Log (infl. + 1) Floral br. length (cm) Log (fl. br. + 1) Sepal length (cm) Log (sep. + 1) Petal length (cm) Log (pet. + 1) Brocchinia reducta Ananas ananasioides Bromelia penguin Broemlia chrysantha Aechmea spectabilis Billbergia macrolepis Cottendorfia florida Guzmania monostachia Tillandsia multicaulis Tillandsia flexuosa Vriesea espinosae Navia ignesicola Pitcairnia maidifolia Pitcairnia meridensis Dyckia ferox Puya alpestris Puya berteroniana Puya weddelliana Puya chilensis Puya gilmartiniae Puya boliviensis Puya castellanosii Puya raimondii Puya retrorsa Puya venusta Puya spathacea Puya pygmea Puya coerulea Puya aequatorialis Puya laxa Puya medica Puya floccosa Puya venezuelana Puya aristeguietae Puya trianae Puya goudotiana Puya ferruginea Puya ferreyrae Puya nitida Puya westii Puya nutans Puya santosii Puya cuatrecasasii 99–100 99–101 100–200 70–150 99–101 98–102 200–400 20–40 25–40 20–150 15–16 16.5–17 128–131 50–60 58–62 120–150 495–505 ? 499–500 149–151 199–200 100–300 950–1200 200–300 90–100 99–100 20–30 199–200 199–200 79–81 20–37 50–200 59–60 299–300 40–200 499–500 250–400 150–200 170–200 249–250 55–70 80–200 99–100 2.00 2.00 2.00–2.30 1.85–2.18 2.00–2.01 2.00–2.01 2.30–2.60 1.32–1.61 1.41–1.61 1.32–2.18 1.20–1.23 1.25–1.26 2.11–2.12 1.71–1.79 1.77–1.79 2.08–2.18 2.70 ? 2.70 2.18 2.30 2.00–2.48 2.98–3.08 2.30–2.48 1.96–2.00 2.00 1.32 2.30 2.30 1.90–1.91 1.32–1.58 1.71–2.30 1.78–1.79 2.48 1.61–2.30 2.70 2.40–2.60 2.18–2.30 2.23–2.30 2.40 1.75–1.85 1.91–2.30 2.00 159–161 160.0 30.0–200.0 80.0–150.0 100.0–140.0 119–120 99–100 16.0–24.0 30.0–40.0 20.0–50.0 ? 39–41 119–120 70.0–130.0 20–60 60–70 99–100 ? 99–100 10.0–20.0 100.0 39–40 123–125 40.0–60.0 30.0–35.0 60.0–100.0 15.0–22.0 59–61 99–100 27–28 20–20.5 100.0–110.0 25.0–34.0 99–100 24.0–28.0 100.0–170.0 99–100 55.0–100.0 35.0–60.0 99–100 15.0–23.5 31.0–37.0 25–27 2.21 2.21 1.49–2.30 1.91–2.18 2.00–2.15 2.08 2.00 1.23–1.40 1.49–1.61 1.32–1.71 ? 1.60–1.62 2.08 1.85–2.12 1.32–1.79 1.79–1.85 2.00 ? 2.00 1.04–1.32 2.00 1.60–1.61 2.09–2.10 1.61–1.79 1.49–1.56 1.79–2.00 1.20–1.36 1.78–1.79 2.00 1.45–1.46 1.32–1.33 2.00–2.05 1.41–1.54 2.00 1.40–1.46 2.00–2.23 2.00 1.75–2.00 1.56–1.79 2.00 1.20–1.39 1.51–1.58 1.43 14.5–15.0 14.5–15.0 30.0–60.0 79.0–81.0 60.0–100.0 39.5–40.0 100.0–300.0 8.0–15.0 14.0–14.5 50–61 6.9–7 4.9–5.0 44–45 14.9–15 ? 99–100 99–100 ? 99–100 50.0–60.0 99–100 39–40 430–500 40.0–80.0 30.0–35.0 40.0–60.0 3.0–5.0 98–101 98–101 70.0–80.0 10.0–20.0 30.0–100.0 18.0–22.0 100.0–110.0 15.0–30.0 100.0–200.0 199–200 60.0–75.0 58.0–62.0 49–50 29–30 32.0–50.0 ? 1.19–1.20 1.19–1.20 1.49–1.79 1.90–1.91 1.79–2.00 161.00 2.00–2.48 0.95–1.20 1.18–1.19 1.78–1.79 0.90 0.77–0.78 1.65–1.66 1.20 ? 2.00 2.00 ? 2.00 1.71–1.79 2.00 1.60–1.61 2.63–2.70 1.61–1.69 1.49–1.56 1.61–1.79 0.60–0.78 2.00–2.01 2.00–2.01 1.85–1.91 1.04–1.32 1.49–2.00 1.28–1.36 2.00–2.05 1.20–1.49 2.00–2.30 2.30 1.79–1.88 1.77–1.80 1.70–1.71 1.48–1.49 1.52–1.71 ? 1.5–2.0 1.5–2.0 1.6–3.0 1.0–1.5 0.1–0.2 3.45–3.5 0.19–0.2 2.3–2.9 4.9–5.0 2.0–3.0 2.4–2.5 2.6–2.7 0.39–0.40 0.7–1.1 0.2–0.3 2.6–3.3 4.4–4.8 1.49–1.5 3.4–5.0 1.5–2.5 2.1–3.3 3.2–3.5 3.1–6.0 3.0–3.5 0.39–0.4 1.3–2.2 2.9–3.0 1.69–1.7 1.79–1.8 1.3–2.0 1.5–1.6 1.3–1.7 4.45–4.5 1.6–1.8 5.0–6.0 2.0–3.7 4.45–4.5 5.0–7.0 3.4–3.8 0.8–1.0 1.99–2.0 3.2–3.3 2.59–2.6 0.40–0.48 0.40–0.48 0.41–0.60 0.30–0.40 0.04–0.04 0.65 0.08 0.52–0.59 0.77–0.78 0.48–0.60 0.53–0.54 0.56–0.57 0.14–0.15 0.23–0.32 0.08–0.11 0.56–0.63 0.73–0.76 0.40 0.64–0.78 0.40–0.54 0.49–0.63 0.62–0.65 0.61–0.85 0.60–0.65 0.14–0.15 0.36–0.51 0.59–0.60 0.43 0.45 0.36–0.48 0.40–0.41 0.36–0.43 0.74 0.41–0.45 0.78–0.85 0.48–0.67 0.74 0.78–0.90 0.64–0.68 0.26–0.30 0.48 0.62–0.63 0.56 0.65–0.7 0.65–0.7 1.2–3.0 0.9–1.7 0.9–2.0 0.99–1.0 0.28–0.3 1.75–1.80 3.5–3.6 2.0–3.0 1.19–1.2 4.4–4.5 2.6–3.0 3.1–3.2 0.5–0.9 2.2–2.9 2.2–2.3 1.79–1.80 3.5–5.8 1.7–2.4 3.19–3.20 2.9–3.0 4.0–4.1 1.99–2.0 1.5–2.0 1.5–2.2 1.5–1.8 2.39–2.40 2.0–2.3 1.69–1.70 1.8–2.0 2.5–3.3 1.8–2.0 2.5–3.3 2.0–2.5 2.0–3.0 1.2–4.5 3.2–3.7 2.9–3.0 1.99–2.0 1.79–1.80 1.3–2.0 2.35–2.4 0.22–0.23 0.22–0.23 0.34–0.60 0.28–0.43 0.28–0.48 0.30 0.11 0.44–0.45 0.65–0.66 0.48–0.60 0.34 0.73–0.74 0.56–0.60 0.61–0.62 0.18–0.28 0.51–0.59 0.51–0.52 0.45 0.65–0.83 0.43–0.53 0.62 0.59–0.60 0.70–0.71 0.48 0.40–0.48 0.40–0.51 0.40–0.45 0.53 0.48–0.52 0.43 0.45–0.48 0.54–0.63 0.45–0.48 0.54–0.63 0.48–0.54 0.48–0.60 0.34–0.74 0.62–0.67 0.59–0.60 0.48 0.45 0.36–0.48 0.53 1.3–1.7 1.3–1.7 1.5–3.0 0.9–1.5 2.4–2.5 4.2–4.3 0.6–1.0 3.0–3.3 6.9–7.0 3.9–4 2.9–3.0 ? 4.9–5.0 6.35–6.40 1.2–1.4 4.8–5.0 5.0–5.6 3.45–3.50 4.9–5.0 3.0–4.8 4.9–5.0 3.9–4.0 7.9–8.0 3.7–4.0 3.4–3.5 2.5–3.3 1.5–2.0 3.7–5.0 2.9–3.0 2.9–3.0 3.4–4.0 3.9–4.0 3.0–3.5 5.0–6.0 3.10–3.20 5.0–6.0 8.0–14.0 3.9–4.0 6.9–7.0 3.9–4.0 3.9–4.0 3.0–3.6 4.90–5.0 0.36–0.43 0.36–0.43 0.40–0.60 0.28–0.40 0.53–0.54 0.72 0.20–0.30 0.60–0.63 0.90 0.69–0.70 0.59–0.60 ? 0.77–0.78 0.87 0.34–0.38 0.76–0.78 0.78–0.82 0.65 0.77–0.78 0.60–0.76 0.77–0.78 0.69–0.70 0.95 0.67–0.70 0.64–0.65 0.54–0.63 0.40–0.48 0.67–0.78 0.59–0.60 0.59–0.60 0.64–0.70 0.69–0.70 0.60–0.65 0.78–0.85 0.61–0.62 0.78–0.85 0.95–1.18 0.69–0.70 0.90 0.69–0.70 0.69–0.70 0.60–0.66 0.77–0.78 C. T. HORNUNG-LEONI and V. SOSA 0 PHYLOGENY OF PUYA SUBGENUS PUYA 101 Figure 3. Strict consensus tree of the nine most parsimonious trees based on discrete characters. Numbers above the branches indicate jackknife support. Numbers below the branches indicate Bremer support [L = 505 steps; consistency index (CI), 0.250; retention index (RI), 0.522]. correlated with petal length (Hornung-Leoni & Sosa, 2005). It is suggested that, in subgenus Puya, large and giant plants have large inflorescences; this is advantageous because, if the flowers are mediumsized, there is an increase in flower number and a larger seed set can be produced. Furthermore, it has been demonstrated that large plants with a large number of flowers are more attractive to pollinators (Kawasaki & Hori, 1999). In the clade of subgenus Puya, the flowers are covered by small to large bracts. It is interesting that P. alpestris and P. berteroniana, species that can be confused, are differentiated by the size of their floral bracts (in the latter species they are larger). Puya chilensis, P. castellanosii, and P. raimondii have flowers with large bracts. More information is needed on the pollination system of the species in this group to understand the evolution of this floral character. It is concluded that, in order to corroborate the infrageneric classification of Puya, an extended sam- pling of the species is needed. Our results clearly show that the species classified in subgenus Puya are retrieved in a group; however, the results do not conclusively determine the taxonomic status of these species. More informative characters would also be useful to resolve the relationships of the species belonging to this large genus. Molecular datasets will undoubtedly provide these variable characters. ACKNOWLEDGEMENTS Edmundo Saavedra kindly prepared the pen and ink illustration. The first author thanks the ‘Red Latinoamericana de Botánica’ for a doctoral scholarship and special funds to travel to Peru. We are grateful for a grant from the International Association for Plant Taxonomy (IAPT) to visit the Chilean herbaria and to the Instituto de Ecología, A.C. for providing support to visit the Colombian herbaria. Thanks are © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 102 C. T. HORNUNG-LEONI and V. SOSA Figure 4. Single most parsimonious tree based on discrete + continuous characters. Numbers above the branches indicate jackknife support. Numbers below the branches indicate Bremer support [L = 530 steps; consistency index (CI), 0.250; retention index (RI), 0.515]. Synapomorphic characters for genus Puya and subgenus Puya are illustrated. due to J. Betancur, J. Manzanares, J. R. Grant, M. Dillon, and B. Holst for their advice, and to G. Montenegro, D. Potojniak, and L. Martínez in Chile, Meri Suni and Giovana Vadillo in Peru, and J. Gaviria, R. Pabón, and J. Bueno in Venezuela, for their help with fieldwork. We are grateful to Ivón Ramírez, Adolfo Espejo, Helga Ochoterena, and two anonymous reviewers for the revision of the manuscript, and also to Anna María Leoni for reviewing a previous version of this paper and Bianca Delfosse for assistance with the English. We thank Pablo Goloboff for his advice on the use of the TNT program. We are grateful to the heads of the following herbaria for access to their collection and the loan of specimens: COL, F, HAL, HDCV, MERC, NY, SGO, US, USM, VEN, and XAL. REFERENCES Archie JW. 1985. Methods for coding variable morphological features for numerical taxonomic analysis. Systematic Zoology 34: 326–345. Baker JG. 1889. Pitcairnia subgenus Puyopsis. Handbook of the Bromeliaceae 91: 117. Barfuss MJJ, Samuel R, Till W, Stuessy TF. 2005. 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Character states, morphological variation and phylogenetic analysis: a review. Systematic Botany 16: 553–583. Terry RG, Brown GK, Olmstead RG. 1997. Examination of subfamily phylogeny in Bromeliaceae using comparative sequencing of plastid locus ndhF. American Journal of Botany 84: 664–670. Thiele K. 1993. The Holy Grail of the perfect character: the cladistic treatment of morphometric data. Cladistics 9: 275– 304. Varadarajan GS, Gilmartin GK. 1988. Morphological variation of some floral features of the subfamily Pitcairnioideae (Bromeliaceae) and their significance in pollination biology. Botanical Gazette 149: 82–91. APPENDIX 1 HERBARIUM SPECIMENS USED FOR THIS STUDY Aechmea spectabilis Brongn. ex Houllet. VENEZUELA: Edo. Trujillo, Entre Trujillo y Boconó, E. Graf s.n. (F). Edo. Mérida, Mpio. Tovar, C. Hornung 209 (MERC). Edo. Mérida, C. Hornung 219 (MERC). Edo. Mérida, Mpio Justo Briceño, P. López 3472 (MERF). COLOMBIA: Dpto. Magdalena, about 11 km south-east of Molino, O. Haught 4051 (F). Ananas ananassoides (Baker) L.B. Sm. BOLIVIA: Santa Cruz, Nuflo de Chavez, Rancho Zapoco, 90 km de Concepcion, T. Killleen 1479 (NY). GUYANE FRANCAISE: Roche Touatou – Basin d’inselberg, J.J. Granville 12988 (NY). BRAZIL: Edo. Amazonia, Municipality if Humaitá, Rio Madeira, road to Humaitá to Labrea, km 20, Savana margin, G.T. Prance 3407 (F). Minas Gerais, Br 153, 20 km S de Frutal, A. Krapocickas 33048 (F). In Cerrado, along Anhuma Creek, 300 km past Cuiaba en route to Goiania, Mato Grosso, B. Maguire (F1769731). Mato Grosso, Pantanal de Paiaguas, Fazenda Buriti, 65–70 km de estrada Cuiaba-Campo Grande, A. Krapovickas 29916 (F). © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 104 C. T. HORNUNG-LEONI and V. SOSA 3–4 km SW of Mutumparaná on railroad to Abuna, G.T.Prance 5453 (F). VENEZUELA: Edo. Bolívar, Basin of Río Parguaza, Croizat 40 (F). Billbergia macrolepis L.B.Sm. BRAZIL: T.F. Do Rondonia, Mpio. De Ariquemes, Mineracao Mibrasa, Setor Alto Candeias, km 128, Sudoeste de Ariquemes, L.O.A.Teixeira 611 (NY). VENEZUELA: Edo. Amazonas, Cerro Sipapo (Paráque), B. Maguire 28053 (NY). Bajo el Río Guaramacal, R. Liesner 10610 (VEN). Brocchinia acuminata L.B.Sm. VENEZUELA: Edo. Territorio Federal Amazonas, Summint to cerro Duida, on high moist ridge top, J.A. Steyermark 58363 (F). Edo. Bolívar, Chimantá Massif, moist forest along quebrada, vicinity of camp 4, south-western edge of Apacará–Tepuí, J. A. Steyermark 75007 (F). Edo. Bolívar, Cerro Guanacoco, cumbre, porción nor-oeste cerca del borde riscoso, J.A. Steyermark 109715 (F). Amazonas, Dpto. Río Negro, Cerro de la Neblina, Camp XI, 6.2 km NNE of Pico Phelps (= Neblina), M. Nee 31098 (NY). Edo. Bolivar, Dto. Piar, summit of Amaruay-tepui, SE quarter of tepui, R. Liesner 20869 (NY). Territorio Federal Amazonas, Sierra Parima, J. Steyermark 107507 (US). Brocchinia reducta Baker. VENEZUELA: Edo. Bolívar, Gran Sabana, Roraima, J. Pruski 1413 (NY). Edo. Bolívar, Dto. Roscio: Valle del Río Aponguao, O. Huber 14.vi.1985 (NY). Edo. Bolívar, Qda. Pacheco, approx. 70 km N de Sta. Elena, N. Xena 313 (NY). Bromelia crysantha Jacq. VENEZUELA: Edo. Mérida, Mpio. Sucre, vía Las Coloradas, C. Hornung 89 (MERC). Edo. Mérida, Mpio. Sucre, San Juna-Las Gonzalez, R. Rico 335 (MERC). Bromelia pinguin L. HONDURAS: Dpto. El Paraíso, Río Choluteca near bridge of Ojo de Agua, L.O. Williams 42888 (NY). MÉXICO: Veracruz, Mozomboa, Mun. Actopan, J. Dorantes 837 (XAL). Veracruz, Chavarillo, 1.5 km delante de la estación de Chavarillo, Mpio. Emiliano Zapata, G. Castillo-Campos 12413 (XAL). Veracruz, La Mancha, Mpio. Actopan, G. CastilloCampos 12339 (XAL). Veracruz, 3 km ak E de Santa Fe, Mpio. Veracruz, V.E. Luna 305 (XAL). Veracruz, Estación biológica El Morro de La Mancha, Mpio. Actopan, B. Guerrero C. 1673 (XAL). Veracruz, 5 km al NO de Panuco, L.I. Nevling 401 (XAL). Veracruz, Carro Topila ejido ‘Benito Juarez’, Mpio. Panuco, C. Gutierrez 1999 (XAL). Veracruz, Ocozotepec, Mpio. Soteapan, H. Cruz Ramírez 11 (XAL). Yucatán, km 4 del crucero rumbo a San Felipe, Mpio. Río Lagartos, E. Ucan 789 (XAL). Yucatán, a 90 km del Crucero, rumbo a Tizimin, Mpio. Río Lagartos, E. Ucan 1155 (XAL), G. Castillo 5762 (XAL). Bahia de Banderas, Mpio. Bahia de Ban- deras, Nayarit (XAL 16364). México, Islas Marietas, Mpio. Bahia de Banderas, Nayarit, G. Castillo 5914 (XAL). PANAMÁ: Prov. Los Santos, between Limon and Punta Mala, T.B. Croat 9746 (NY). Cottendorfia florida Schult. f. BRAZIL: Mpio. De Macujê, 10–17 km ao NW de Mucujê, na estrada para Andaraí, S.A. Mori (US 2854490); Mpio. De Macujê, 10–17 km ao NW de Mucujê, na estrada para Andaraí, S.A. Mori (US 2854493). Bahia, Mun. Rio de Contas Pico das Almas, Vertente leste, Na parte norte do vale abaixo do pico, R.M. Harley 26196 (F). South of Andarí, along road to Mucugé near small town of Xique-Xique, R.M. Harley 18687 (NY). Río Apiaba (mun. Mucugê), Bahia, G. Hatschbach 47534 (NY). Dyckia ferox Mez. ARGENTINA: Dpto. Capital, Prov. Corrientes, Procedente de San Roque, A. Schinini 13850 (F). Dpto. Capital, Ruta 12 y Ayo, Richuelo, A. Schinini 12439 (F). Guzmania monostachya Rusby ex Mez. COSTA RICA: H.E. Stork 2933 (F). NICARAGUA: G.S. Bunting 352 (F). La Selva, U. Rowlatt 1851 (F). ECUADOR: Prov. Carchi, environs of Chical, 12 km below Maldonado on the río San Juan, M.T. Madison 4484 (F). Dpto. Tumbez, Prov. Tumbez, mts E of Hacienda Chicama, A. Weberbauer 7643 (F). PERÚ: Dpto. Cajamarca, Prov. Santa Cruz, alrededores de Monteseco, A. Sagástegui 12362 (F). Dpto Cajamarca, Prov. Santa Cruz, Distr. Catache, Upper Río Zaña valley, M. Dillon 4371 (F). (L.Rusby) ex Mez; Dpto. San Martin, San Roque, L. Williams 7238 (F). VENEZUELA: Edo. Mérida, Mpio. Libertador, C. Hornung 210 (MERC). Edo. Mérida, Mpio. Libertador, C. Hornung 236 (MERC). Edo. Mérida, Mpio Libertador, Campo de Oro, T. Ruiz 14662 (MERF). Edo. Mérida, Mpio. Libertador, Campo de Oro, T. Ruiz 14656 (MERF). USA: Florida, s.c, s.n (F 167005). Prov. De Heredia, Finca La Selva, the OTS Field Station on the Río Puerto Viejo just E of its junction with the Río Sarapiquí, T. McDowell 353 (F). USA: Subtropical Florida, N.L. Buten 257 (F). Navia igneosicola L.B.Sm., Steyerm. & H.Rob. VENTerritorio Federal Amazonas, Dpto. Atures: forested areas and igneous outcrops along Río Coromoto, at Tobogán de la Selva, J. Steyermark 122478 (US). EZUELA: Pitcairnia maidifolia Decne. ex Planch. COLOMBIA: Dpto Antioquia, Mpio. De Cocorná (Ant.), camino entre ‘la Piñuela’ y ‘La Vega’, margen izquierda del Río Santo Domingo, R. Fonnegra s.n. (F 2181034); Mpio. Amalfi, Verede El Oso, Cordillera Central, J. Betancur 916 (F). COSTA RICA: San José, Dota, Zona Protectora Cerro Nara, Faldas del Cerro Nara, © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 PHYLOGENY OF PUYA SUBGENUS PUYA O. Valverde 515 (F). Prov. San José, F. Montgomery G. 5052 (F). GUIANAS: Region Cuyuni-Mazaruni, Mt. Holitipu, below peak, T. McDowell 3027 (F). HONDURAS: Dpto. Morazán, Agua Amarilla, L.O. Williams 10814 (F). Dpto. Morazán, Region of Agua Amarilla, above El Zamorano, P.C. Standley 5061 (F). Dpto. Morazán, north-west of El Zamorano, P.C. Standley 22890 (F). Dpto. Morazán, along and near Río Agua Amarilla, above El Zamorano, P.C. Standley 12813 (F). El Paraiso, 3 km north of Las Manos, L.O.Williams 42265 (F). Dpto. Morazán, Region of Agua Amarilla, above El Zamorano in pine–oak forest, P.C. Standley 10 (F). VENEZUELA: Edo. Bolivar, Massif of Chimantá, J.A. Steyermark 75475 (F). Territorio Federal Amazonas, Sierra Parima, J.A. Steyermark 105910 (F). Territorio Federal Amazonas, Cero Duida. J.A. Steyermark 57969 (F). Edo. Mérida, between El Molino and ridge above San Isidro, J.A. Steyermark 56519 (F). Edo. Mérida, Mpio. Libertador, C. Hornung (MERC). Edo. Mérida, Mpio. Cardenal Quintero, C. Hornung 136 (MERC). Edo. Mérida, Mpio.Tovar, C. Hornung 174 (MERC). Edo. Mérida, Mpio. Tovar, C. Hornung 175 (MERC). Edo. Mérida, Mpio. Tovar, C. Hornung 176 (MERC). Edo. Mérida, Mpio. Tovar, via Zea, C. Hornung 177 (MERC). Edo. Mérida, Mpio. Arzobispo Chacón, C. Hornung 196 (MERC). Edo. Mérida, C. Hornung 221 (MERC). Edo. Mérida, Mpio. Tovar, Zea, P. López 1236 (MERF). Edo. Mérida, Dtto. Justo Briceño, López 3513 (MER, MERF). Pitcairnia meridensis Klotzsch ex Mez. VENEZUELA: Edo. Mérida, between El Molino and ridge above San Isidro, J.A. Steyermark 56519 (F). Edo. Mérida, Mpio. Campo Elías, L. Aristeguieta 2485 (VEN); Edo. Mérida, Ca. Jají, F. Oliva 263 (VEN). Edo. Mérida, Mpio. Rivas Dávila, B. Marcano 1743 (VEN). Edo. Mérida, Mpio. Rivas Dávila, C. Benitez 2072 (VEN). Edo. Mérida, J.A. Steyermark 56519 (VEN). Edo. Mérida, Mpio. Campo Elías, F. Oliva 228 (VEN). Edo. Mérida, Carbonera-Azulita, J. de Brujin 1131 (VEN). Mpio. Dávila, B. Marcano 1743 (MER). Edo. Mérida, Estación teleférico La Montaña, D.L. Kelly 9099 (MER). Edo. Mérida, Mpio. Rivas Dávila, P.S. López 10 (MERF). Edo. Mérida, Mpio. Libertador, T. Ruiz 8741 (MERF). Edo. Mérida, Mpio. Sucre, T. Ruiz 811 (MERF). Edo. Mérida, Mpio. Libertador, C. Hornung 74 (MERC). Edo. Mérida, Minas de Bailadores, C. Hornung 107 (MERC). Edo. Mérida, Mpio. Andrés Bello, C. Hornung 143 (MERC). Edo. Mérida, Mpio. Arzobisco Chacón, C. Hornung 194 (MERC). Edo. Mérida, Mpio. Arzobispo Chacón, C. Hornung 198 (MERC). Edo. Mérida, C. Hornung 216 (MERC). Puya aequatorialis Ed. André. ECUADOR: Azuay, Prov. Azuay, between ríos Azogues and Gualaceo, valley of the río Paute, between Puate and Cuenca, W.H. Camp 105 2322 (US). A 15 km de Quito, Otavalvo-70 m terreno volcanico, A. J. Gilmartin 1084 (US). André 3594 (F). Loja, Cuenca-Ona-Zaraguro, M.B. Foster 2608 (F). Azuay, valley of the río Paute, between Paute and Cuenca, W.H. Camp 3222 (US). Vicinity of Cumbe, J.N. Rose 22954 (NY). Prov. Azuay, valley of the río Paute, between Paute and Cuenca, W.H. Camp 2322 (NY). Puya alpestris (Poepp.) Gay. CHILE: 1877. SGO (046422); J. West 4981 (US); F (739422); F (1670523). N. Floy Bracelin 2802 (F). N. Floy Bracelin 2766 (F). La Hermida, cerca de Santiago, G. Looser 2104 (F). Cordillera de Colchagua (SGO 046423). Cerro Sur Baños Flaco, M. L. Espinosa (leg) s.n. xii.1937 (SGO). Coordillera de Colchagua, s.n. (SGO 046425). Reiche s.n. dic/91 (SGO 061082). Fundo Fray Jorge, SGO (060188); s.n. V/1884 (US photo). Prov. Concepción, Hills north to Concepción, J. West 4981 (US). Cerro San Cristóbal, Germain s.n., 1854 (SGO). Coquimbo, M. Muñoz 909 (SGO). Coquimbo, C. Muñoz 1552 (SGO). Coquimbo, C. Muñoz 1302 (SGO). Cerro Fray Jorge, Philippi s.n. (SGO). Fray Jorge, Ovalle, Philippi s.n., I/1883 (SGO). Cordillera de La Dehesa, Philippi s.n., XI/1861 (SGO). Región metropolitana, Carr. Santiago-Farellones, C. Hornung 1109 (HDCV). Puya aristeguietae L.B.Sm. VENEZUELA: Edo. Trujillo, Dto. Boconó, Páramo de Guaramacal, L.J. Dorr 5000 (NY). Edo. Trujillo, Alrededores de Guirigay, L. Aristeguieta 3539 (NY). Edo. Trujillo, Dto. Boconó, Páramo de Guaramacal, SE of Boconó, L.J. Dorr 7326 (NY). Edo. Mérida, Bosque exp. ‘San Eusebio’, Dpto. Campo Elías, J.P. Schulz 388 (NY). Edo. Mérida, Dtto. Rangel, Quebrada ‘Puya’ (unnamed Qba.) c. 3–4 km S of the mouth of the Rio Los Granates, Parque Nacional Sierra Nevada, L.J. Dorr 5598 (NY). Edo. Trujillo, Páramo de Guirigay, L. Aristeguieta 3539 (US). Edo. Mérida, Mpio. Rangel, C. Hornung 161 (MERC). Edo. Mérida, Mpio. Páramo Gaviria, C. Hornung 208 (MERC). Edo. Mérida, Mpio. Rangel, T. Ruiz 7310 (MERF). Edo. Mérida, Mpio. Miranda, T. Ruiz 12267 (MERF). Edo. Mérida, Mpio. Miranda, T. Ruiz 12268 (MERF). Edo. Mérida, Parque Nacional Sierra Nevada, A.P. Yañez 1719 (MER). Edo. Trujillo, Páramo Guirigay, L. Aristeguieta 3539 (VEN). Puya berteroniana Mez. BOLIVIA(?): near La Paz (probably the label is an error), Rusby 2850 (US). CHILE: Prov. Curicó, Hacienda Monte Grande, E. Wedermann 563 (F). IIX Región, Prov. Santiago, Peñaflor cerro, G. Montero 768 (F). CHILE: Limache, Belen s.n. 10.ii.1917 (F). Rancagua, Bertero 115 (F photo). Viña del Mar, 16.xii.1923 (F). Cuesta de La Dorminda, L. Gonzalez s.n. 26.v.1983 (HDCV). Cuesta La Dormida, bajando del Roble, C. Hornung 1111 (HDCV). Haci- © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 106 C. T. HORNUNG-LEONI and V. SOSA enda Rinconada La Cesda, Maipu, Qda La Plata, F. Schlegel 1664 (SGO). Prov. Santiago, Región Metropolitana, Reserva Forestal Río Clarillo, J. Yañez s.n. 3.viii.1993 (SGO). Prov. Valparaiso, four km southwest of Valparaiso, P.C. Hutchison 22 (SGO). Prov. Coquimbo, IV Región, Coquimbo, M. Muñoz S. 908 (SGO). IV Región, Playa Las Tacas, 20 km al Sur de Coquimbo, A.R. Flores s.n. 15.viii.1986 (SGO). Rancagua, Bertero 115 (US). Valparaiso, Weber 3252 (US). Concón, Miers 347 (US). Puya boliviensis Baker. BOLIVIA (now Chile): Cobija, Gaudichau, Julliet s.n. 1836–1837 (F 1435100). Cobija, Gaudichaud s.n. (F 741178). Gaudichaud s.n. (US 2144992). Gaudichau s.n. (F1435081). CHILE: Antofagasta región II, Prov. Antofagasta, Cero Perales, c. 5 km E of Taltal, M.O. Dillon 5377 (F). Quebrada Paposo, c. 5–7 km E of Caleta, M.O. Dillon 5242 (F). CHILE: Antofagasta región II, Prov. Antofagasta, Cerro Perales, c. 5 km E of Taltal, M.O. Dillon 5377 (F). II Región, Qda. Rinconada de Paposo, A. Hoffmann s.n. 2.xii.1988 (SGO). CHILE: II Región, Mirador de Paposo, subiendo por quebrada Los Yales, A. Hoffmann s.n. 1.xii.1988 (SGO). II Región, qda. en camino a Cifuncho, A. Hoffmann s.n. 1.xii.1988 (SGO). II Región, Qda. Matancilla, A. Hoffmann s.n. 3.xii.1988 (SGO). II Región, Taltal, Qda. Matancilla, A. Hoffmann s.n. 3.xii.1988 (SGO). Morro de Caldera (SGO), Paposo, Reich 360 (SGO). Gaudichaud s.n. (US 21449992). Region II, Antofagasta, Prov. Antofagasta, Cerro Perales, c. 5 km E of Taltal, M.O. Dillon 5377 (F). Puya castellanosii L.B.Sm. ARGENTINA: Cachi, Prov. Salta, Dpto. Chachi: Brealito, G.S. Varadarajan 1476 (US). Dpto. Cachi, Prov. Salta, Brealito, G.S. Varadarajan 1476 (US). Prov. Salta, Catellanos s.n. [type], 1897 (US). Dpto. Molinos, Prov. Salta, Brealito, T. Meyer 9164 (US). Salta, Laguna del Brealito (Valle Calchaqui), Castellanos 45819 (US). Puya chilensis Molina. BOLIVIA (now CHILE): Gaudichaud, s.n. (F 1370754). CHILE: Limache, Frisco (F 633928). 1837 (F 1435147). Viña del Mar, 7.ix.1922 (F). Viña del Mar, 7.ix.1922 (F). Prov. Elqui, T.G. Lammers, C.M. Baeza P. & P. Peñalillo B. 7653 (F). Angol, Philippi s.n., i.1877 (SGO). Constitución, K. Reiche s.n., xii.1891 (SGO). Fundo Fray Jorge, C. Carrizo s.n., 10.iii.1947 (SGO). Cordillera de Colchagua, L. Landbeck s.n., xii.1860 (SGO). Cordillera de Colchagua, Philippi s.n. (SGO 6425). Cerro, Sur Baños Flacos, M. Espinosa s.n., xii.1937 (SGO). Zapallar, Philippi s.n., ix.1875 (SGO). Aconcagua: Zapallar, Philippi s.n., ix.1865 (SGO). Paposo, K. Reiche s.n., ix.1909 (SGO). Cordillera de Cauquenes, K. Reiche s.n., x.1907 (SGO). Angol, Philippi s.n., 1877 (SGO). Constitución, K. Reiche s.n., xii.1891 (SGO). Chillan, Philippi s.n., xii.1869 (SGO). Llico, Philippi s.n., xii.1861 (SGO). Puya coerulea Miers. CHILE: Angostura de Praine, G. Looser s.n., 4.xii.1932 (F). s.n. (F1668607), Angostura de Praine, G. Looser 2553 (F). Angostura de Praine, Looser 2550 (F). Angostura de Praine, G. Looser 2549 (F). Angostura de Praine, G. Looser 2551 (F). Prov. Santiago, P.C. Hutchinson 202 (F). Phillipii s.n. (F 741265). Prov. Curicó: Hacienda Monte Grande, E. Wedermann 539 (F). s.n. (F 835815). Río Clarillo, s.n. (HDCV 949), Vía El Roble, C. Hornung 1110 (HDCV). Baños de Cauquenes, Philippi s.n., iii.1875 (SGO). Llico, Philippi s.n., xii.1861 (SGO). Inter Poblacion El Cueva, Philippi s.n. (SGO 46407). Chillan, Philippi s.n., xii.1869 (SGO). Cordillera de Popeta, Philippi s.n., i.1884 (SGO 46413). Angol, Philippi s.n., i.1877 (SGO). Complejo Turistico La Leonera, M. Muñoz & S. Moreira 2393 (SGO). Complejo Turistico La Leonera, M. Muñoz & S. Moreira 2394 (SGO). Las Tacas, A. Flores s.n., 15.viii.1986 (SGO). Taltal, A. Hoffmann & A. Flores s.n., 3.xii.1988 (SGO). Quebrada Camino a Cifuncho, A. Hoffmann & A. Flores s.n., 1.xii.1988 (SGO). Puya cuatrecasasii L.B.Sm. COLOMBIA: Dpto. Valle, Cordillera Central: entre Pan de Azúcar, J. Cuatrecasas 27573 (COL). Cordillera Central, vertiente occidental, J. Cuatrecasas 18962 (COL). Dpto. Cauca, Macizo Colombiano, Páramo de las Papas, J. Idrobo 4062 (COL). Dpto. del Cauca, Cordillera Central, vertiente occidental, Cabeceras del Río Palo. J. Cuatrecasas 18962 (F). Tolima, above ‘Ampilio’, Cabeceras río Ereje, E. L. Core s.n. 21.xi.1944 (F). Puya ferreyrae L.B.Sm. ECUADOR: Prov. Loja: Las Chinchas, Reg. Central, M. Acosta S. 7803 (F). PERU: Dpto. La Libertad, Prov. Otuzco, road to Huamachuco, 11 km above Samne, P. C. Hutchinson 6124 (F). Dpto Cajamarca, Sangal (San Pablo), Prov. San Pablo, A. Sagástegui 15367 (F). PERU: Dpto. Cajamarca, c. 7 km E of Magdalena, M.O. Dillon 6206 (F). Dpto. Cajamarca, Prov. Cajamarca. Entre Magdalena – Chilete, en la parte alta de la hacienda la Viña, I. Sanchez V. 3574 (F). Dpto. Cajamarca, Prov. Cajamarca, Dist. Jesús, a 1 km de la localidad de Jesús, siguiendo la carretera, I. Sanchez V. 6153 (F). Dpto. La Libertad, Prov. Otuzco, Casmiche (Ssmne-Otuzco). A. Sagastegui 11515 (F). Road to Huamachuco, 11 km above Samne, J. Kenneth 6124 (F). Puya ferruginea (Ruiz & Pav.) L.B.Sm. BOLIVIA: Nequejahuira, SC 634, 15.v.1926 (NY). Dpto. La Paz, Prov. Murillo, 1 km SW of Mallasilla Golf Course, 9 km SSE of centre of La Paz, M. Nee 34161 (NY). Dpto. La Paz, Province of Murillo, J.C. Solomon 6668 © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 PHYLOGENY OF PUYA SUBGENUS PUYA (NY). Dpto. La Paz, Prov. Murillo, c. 0.4 km NE of Alto Irpavi, Solomon J.C. 6093 (NY). O. Kuntze s.n. 13–21.iv.1892 (NY). La Paz, s.n. 152, 1890 (NY). H.H. Rusby 2845, iv.1885 (NY). Yungas, H.H. Rusby 2847, 1885 (NY). Dpto. La Paz, Prov. Nor Yungas, 3.6 km NE of (below) Chupipata on road to Yolosa, J.J. Solomon 15642 (NY). Dpto. La Paz, Prov. Nor Yungas, 4.7 km al NE (abajo) de Chuspipata, J.C. Solomon 17336 (NY). Dpto. La Paz, Prov. Murillo, M. Nee 34161 (NY). Nequejahuira, G.H. Tate 634, 15.v.1926 (NY). Yungas, H.H. Rusby 2847 (F). Dpto. La Paz, O. Buchtien s.n. iii.1913 (F). Dpto. La Paz, O. Buchtien, iii.1923 (F). Dpto. La Paz, 1890 (F 162549). Dpto. La Paz, O. Buchtien s.n. iii.1913 (NY). Prov. Huaura, Lomas de Lachay, A. Cano 7081 (USM). ECUADOR: Prov. Loja, Valle Seco de Playas, Catacocha, M. Costa Solís 7998 (F). E. Andre 4019 (NY). PERU: Dpto. Junin, Tarma, E.P. Killip 21806 (NY). G. Mandon 1173 (NY). Yungas, H.H. Rusby 2847 (NY). La Paz, R.S. Williams 2355 (NY). Cordillera Real, Nequejahuira, H.H. Tate 634 (NY). Dpto. Cusco, Prov. Calca, Pisaq ruins, J.D. Boeke 1532 (NY). Dpto. Lima, Prov. Huarochiri, Rio Blanco, P.C. Hutchinson 573 (NY). Dpto. Cusco, Prov. Calca, Lares Valley above Mantoc, A. Weberbauer 7915 (NY). Cordillera Yanachaga, A. Gentry 35919 (NY). Dpto Ancash, Prov. Huaylas, Pueblo Libre, A. Cano 8882 (USM). Dpto. Lima, Prov. Huarochirí, Sangallaya, E. Cerrate 1775 (USM). Dpto. Lima, Prov. Huarochirí, Sangallaya, R. Ferreyra 9200 (USM). Pasco, Cordillera Yanachaga, A. Gentry 35919 (USM). Prov. Huaura, Lomas de Lachay, A. Cano 7081 (USM). Dpto. Cuzco, Prov. Urubamba, E.W. Davis 1774 (USM). Dpto. Cuzco, Prov. Urubamba, W. Davis 1488 (USM). Cuzco, Prov. Urubamba, Sist. Huayllabamba, A.Tupayachi 931 (NY). Dpto. La Libertad, Prov. Trujillo, Cerro Campana. A. Sagástegui 12957 (F). Dpto Lima, Prov. Huarochiri, Río Blanco, Canyon of the Río Rimac, P.C. Hutchinson 573 (F). Dpto. La Libertad, Prov. Trujillo, A. Sagástegui 10980a (F); J. Francis Macbride 3148 (F); Dpto. Yucay, J. Soukup, xii.1937 (F); Dpto. Cuzco, Prov. Urubamba, Chicon Canyon, on rocky slopes, C. Vargas 11091 (F); Cuzco, Colinas de Sacsahuaman, J. Soukup 42 (F). Río Blanco, J.F. Macbridge 3005 (F). Dpto. Lima, Prov. Huarochiri, Infiernillo, T.H. Goodspeed 110601 (F). Dpto. Cuzco, Prov. Urubamba, Chincheros, E.W. Davis 1774 (F). Yanano, J. Francis Macbride 3812 (F). Río Blanco, J. Francis Macbride 711 (F). Dpto. La Libertad, Prov. Otuzco, Cascasday (CollambaySinsicap), A. Sagástegui 15638 (F). Lima, Prov. Huarochiri, Río Blanco, canyon of the río Rimac, P.C. Hutchinson 573 (F). Cusco, Prov. Paucartambo, valle del Pilcopata, road from Patria to Pillahuata, R. Foster & T. Watcher 7482 (F). Prov. Yanachaga, A. Gentry 35919 (F). Dto. Huayllabamba, Cuzco: Prov. Urubamba, A. Tupayachi 931 (NY). Prov. Huarochiri, 107 Río Blanco, canyon of the río Rimac, P.C. Hutchinson 573 (F). Puya floccosa E. Morren. BRAZIL: Amazonia, Territorio do Roraima, G.T. Prance 4532 (NY). Serra Tepequem, terr. Do Rio Branco, B. Maguire 40039 (NY). COLOMBIA: Dpto. Cundinamarca, Cordillera Oriental, Chuneca Creek, 5 km from Ubalá, M.L. Grant 10189 (NY). Dpto. Huila, Cordillera Oriental, east of Neiva, H.H. Rusby 1122 (NY). Dpto. Santander, northern slope of Mesa de los Santos, Eastern Cordillera, E.P. Killip 15002 (NY). Dpto. Cundinamarca, Cordillera Oriental, vertiente Magdalenense, J. Betancur 3975 (COL). VENEZUELA: Edo. Barinas, San Isidro, 27 km de Barinitas, Dtto Bolivar, G. Aymard 2190 (NY). Edo. Bolívar, Dtto. Roscio, sabanas en el valle del Río Kukenán inferior, en la región de Campo Alegre, a aprox. 14 km al SW de S. Ignacio de Yuruaní, O. Hubber 7592 (NY). Edo. Táchira, Dtto. Uribante, S base of Cerro El Morro, J.L. Dorr 7086 (NY). Edo. Bolívar, 17 km E of Canaima, G.T. Prance 28461 (NY). Edo. Bolívar, 3 km S of El Paují, R.L. Liesner 19875 (NY). Edo. Bolivar, Uaipan-tepui, plateau at SE foot of the peak of Uaipan, exposed sandstone shield, T. Koyama 7371 (NY). Edo. Bolívar, Dtto. Rosci, O. Huber 9201 (NY). Edo. Mérida, C. Hornung 206 (MERC). Edo. Mérida, C. Hornung 81 (MERC). Puya gilmartiniae G.S.Varadarajan & A.R.Flores. CHILE: Region IV, Coquimbo, Prov. Elqui, Punta Arrayán, c. 20 km N of La Serena, M.O. Dillon 5449 (F). Region IV, El Olivar, N de La Serena, A. Flores s.n. 16.viii.1986 (SGO). Puya goudotiana Mez. COLOMBIA: Dpto. Cundinamarca, Macizo de Bogotá, cerro Diego Largo, vert. E., J. Cuatrecasas 5162 (F). Dpto. Cundinamarca, Eastern Cordillera, municipality of Calera, Hacienda La Siberia, Páramo de Palacio, R. Merril K. 6039 (F). Dpto. Cundinamarca, Cordillera Oriental, Chocontá, J. Cuatrecasas 9660 (F). Dpto. Cundinamarca, Cordillera Oriental, Páramo de Guasca, J. Cuatrecasas 13547 (F). Dpto. Norte de Santander, Cordillera Oriental, J. Cuatrecasas 10039 (F). Dpto. Cundinamarca, Cordillera Oriental, Macizo de Bogotá, J. Cuatrecasas 7969 (F). Dpto. Cundinamarca, Eastern Cordillera, Municipality of Calera, Hacienda La Siberia, Páramo de Palacio, R. Merrill K. 6039 (F). Dpto. Cundinamarca, Cordillera Oriental, vertiente oriental, Páramo de Guasca, J. Cuatrecasas 9506 (F). Dpto. Cundinamarca, Cordillera Oriental, Páramo de Guasca, vertiente oriental, J. Cuatrecasas 13547 (F). Dpto. Cundinamarca, Cordillera Oriental, Páramo de Chocontá, J. Cuatrecasas 9660 (COL). Dpto. Amazonas, Páramo de Guasca, A. Fernández P. 5760 © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 108 C. T. HORNUNG-LEONI and V. SOSA (COL). Dpto. Cundinamarca, Mpio. La Calera, C. García R. 85 (COL). Dpto. Cundinamarca, Eastern Cordillera, Municipality of Calera, Hacienda La Siberia, Páramo de Palacio, R. Merrill K. 6039 (NY). G. Gutierrez 247 (NY). Bogotá 26.xi.1852 (NY). Dpto. Santander, Vicinity of Charta, E.P. Killip 18930 (NY). Puya laxa L.B.Sm. BOLIVIA: About Pulquina, Santa Cruz, M. Cardenas 5092 (US). About Pulquina, Santa Cruz, M. Cardenas 5092 (US). Puya medica L.B.Sm. PERU: Shorey, Prov. Santiago de Chuco, Sagástegui, Aldave, Fernandez & Fukushima 6175 (XAL). Puya nitida Mez. COLOMBIA: Dpto. Cundinamarca, Cordillera Oriental, Páramo de Guasca, J. Cuatrecasas 13539 (F). Macizo de Bogotá, Cerro entre quebrada de Las Delicias y la de Las Ninfas, J. Cuatrecasas 5629 (F). Dpto. Cundinamarca, Coodillera Oriental, Páramo de Zipaquirá, J. Cuatrecasas 9529 (F). Macizo de Bogotá: Cerro de Guadalupe, J. Cuatrecasas 7958 (F). Dpto. Cundinamarca, Mpio. Guatavita, Laguna de Guatavita, Páramo de Guatavita, J. Betancur 2713 (COL, NY). Dpto. Cundinamarca, Páramo de Cruz Verde, Cordillera Oriental, vertiente occidental, G. Gutierrez V. 366 (F). Boyaca, Paramo de Guina, Sta. Rosita, A.M. Cleff 9740 (NY). M.J. Goudot, 1844 (NY). Cundinamarca, Macizo de Bogotá, Mpio. De Calera, Páramo del Palacio, R.E. Schultes 18727 (NY). Dpto. Boyacá, Páramo de Guina, Sta. Rosita, A. M. Cleef 9740 (COL). Puya nutans L.B.Sm. ECUADOR: Azuay, Campo, W.H. E. 2291 (NY, US). PERU: Cuzco, Cano, A. 4324 (F). Puya pygmaea L.B.Sm. BOLIVIA: Lchmamb 589 (F). ECUADOR: Prov. Azuay, Páramo de Tinajillas, W.H. Camp 2236 (F, US). Prov. Morona-Santiago, Páramo de Castillo, G.S. Varadarajan 1433 (US). Prov. Azuay, Páramo de Tinajillas, S.E. Clemants 2199 (US). PERU: Dpto. Cusco, Prov. Paucartambo, Tres Cruces, Parque Nacional Manu, A. Cano 4563 (F). Prov. Paucartambo, Dpto. Cusco, Tres Cruces P.N.M, A. Cano 3455 (USM). Dpto. Cusco, Prov. Paucartambo, Altura de Teleban P.N.M., A. Cano 3782 (USM). Dpto. Cusco, Prov. Paucartambo, Tres Cruces P.N.Manu, A. Cano 4563 (USM). Dpto. Cusco, Prov. Paucartambo, Acjanaco, PN Manu, A. Cano 3393 (USM). In this study, we extend the distribution of this species to Bolivia and Peru, following the determination from H. Luther, A. Cano, and L. B. Smith, and confirm the characteristics specific to this species (Manzanares, 2005) as the presence of floral bracts erect (vs. reflexes), amongst others. Puya raimondii Harms. BOLIVIA: Huacanqui, M. Cárdenas 4380 (US). Comanche, M.B. Foster 2566 (US). Cochabamba, M.B. Foster 2546 (US). Dpto. Cochabamba, Prov. Arani, G. Schmitt & D. Schmitt 84 (US). La Paz, Prov. Pacajes, Comanche, J.N. Rose & Mrs Rose 18875 (US). La Paz, Pacajes, Comanche, J. Luteyn, L. Dorr, D. Smith & M. Buddensick 13840 (US). Dpto. La Paz, Prov. Pacajes, Comanche, St.G. Beck 2353 (US). PERÚ: La Libertad, Prov. Otuzco, A. Sagástegui, S. Leiva & C. Tellez 14510 (F). Weberbauer 2955 (F). La Libertad, Prov. Otuzco, S. Leiva, P. Leiva & E. Zavaleta 292 (F). Estation 30 miles from Huaraz, Pomopampa, Macbride & Featherstone s.n, 4.xii.1922 (F). Dpto. Puno, Prov. Melgar, H. Ilties & Don Ugent 1288 (US). Prov. Bolognesi, Huishcashpampa, E. Cerrate 2072 (USM). Prov. Huaylas, Dpto. Ancash, A. Cano 6405 (USM). 1/2 km SE of Hacienda Santa Rosa de Achaco, s.n. (USM 159980). Dpto. Huancavelica, carretera CastrovirreynaAyacucho, s.n. (USM 159979). Dpto. Ancash, P.N. Huascarán, C. Hornung 1118 (USM). Dpto. Ancash, Canchayllo, C. Hornung 1120 (USM). Dpto. Ancash, P.N. Huascarán, C. Hornung 1121 (USM). Dpto. Ancash, P.N. Huascarán, C. Hornung 1122 (USM). Dpto. Ancash, P.N. Huascarán, Pumapashimin, C. Hornung 1123 (USM). Dpto. Ancash, Cordillera Negra, C. Hornung 1124 (USM). Dpto. Ancash, C. Hornung 1126 (USM). Puya retrorsa A.J.Gilmartin. ECUADOR: Prov. Chimborazo, between Cajabamba and Pallatanga, G.S. Varadarajan 1440 (US). Prov. Pichincha, Paramo de Huamani, G.S. Varadarajan 1420 (US). Tunguragua, A.J. Gilmartin 1103 (US). Puya santosii Cuatrec. COLOMBIA: Dpto. Cundinamarca, Cordillera Oriental, Páramo de Cruz Verde, J. Cuatrecasas 9518 (F). Dpto. de Cundinamarca, Macizo de Bogotá, Páramo de Usaquén, J. Cuatrecasas 9441 (F). Dpto de Cundinamarca, Páramo de Usaquén, J. Cuatrecasas 7996 (F). Dpto. de Cundinamarca, Páramo de Cruz Verde, Cordillera Oriental, J. Cuatrecasas 10468 (F). Dpto. Cundinamarca, Macizo de Bogotá, Páramo de Uraquén, J. Cuatrecasas 9441 (COL). Meta: Páramo de Sumapáz, Hoya El Nevado, A. Cleef 1503 (COL). Dpto. Cundinamarca, Cordillera Oriental, Páramo de Cruz Verde, J. Cuatrecasas 10468 (COL). Dpto. Cundinamarca, Cordillera Oriental, extremo sudeste de la Sabana de Bogotá, en San Miguel, J. Cuatrecasas 12042 (F). Dpto. Cundinamarca, Cordillera Oriental, Páramo de Cruz Verde, G. Gutierrez 386 (F). Dpto. Cundinamarca, Cordillera Oriental, J. Cuatrecasas 9518 (F). Puya spathacea (Griseb.) Mez. ARGENTINA: Prov. De Cordoba, Sierra Grande, A.P. Rodrigo 458 (NY). © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 PHYLOGENY OF PUYA SUBGENUS PUYA 109 Jujuy, O. Kuntze s.n. x.1892 (NY). Prov. de Córdoba, Sierra chica, G. Hieronymus, s.n. 9.viii.1876 (NY). O. Kuntze, s.n. x.1892 (F). Prov. Córdoba, La Cumbre, A. Haurteig 203 (F). Jujuy, O. Kuntze x.1892 (NY). La Rioja, Guanchín, Castellanos 27/1902 (NY). Prov. Cordoba, Las Cumbres, A. Haurteig 263 (F). BOLIVIA: Troll 661 (F). ARGENTINA: Jujuy, O. Kuntze s.n. x.1892 (F). Hieronymus s.n. (F). Prov. de Cordoba, Loc. La Falda, Cerro El Chorrito, M.M. Job 453 (F). Jujuy, O. Kuntze s.n. ix.1892 (NY). Puya weddelliana Mez. BOLIVIA: Dpto. Chuquisaca, D.S. Correll 644 (US). Tarija, Weddell 4001 (US photo). Puya trianae Baker. COLOMBIA: Dpto. Cundinamarca, Macizo de Bogotá, Páramo de Chisacá, Laguna Negra, J. Cuatrecasas 25924 (COL, F). Dpto. Cundinamarca, Páramo de Chisacá, T.R. Soderstrom 1286 (F). Dpto. Cundinamarca, Andes de Bogotá, Páramo de Cruz Verde, s.n. (COL). Dpto. Cundinamarca, Páramo de Chisacá, T.R. Soderstrom 1286 (F). Dpto. Cundinamarca, Páramo de Cruz Verde, Cordillera Oriental. J. Cuatrecasas 10474 (F). Dpto. Antioquia, Mpio. Urrao, Páramo de Frontino, J. Betancur 1166 (F). Dpto. Cundinamarca, Páramo de Chisacá, en pendientes y colinas, J.M. Idrobo 6525 (F). Cundinamarca, Triana 1314 (F). Dpto. Cundinamarca, Macizo de Bogotá, Páramo de Chisacá, J. Cuatrecasas 25745 (NY). Dpto. Cundinamarca, Macizo de Sumapaz, J. Cuatrecasas 27030 (NY). Dpto. Boyacá, Páramo de la Rusia, Serranía Peña Negra, A.M. Cleef 7430 (COL). VENEZUELA: Edo. Mérida, Mpio. Campo Elías, T. Ruiz 8000 (MERF). Tillandsia flexuosa Sw. COLOMBIA: Vaupes, Río Kuduyarí, Yapobodá, sandstone savannah near headwaters, R.E. Shultes 18487 (F). PANAMÁ: Prov. Cocle, s.n. (F 1693109). Prov. Panamá, Las Sabanas, s.n. (F 686341). VENEZUELA: Mpio Sucre, Edo. Mérida, C. Hornung 48 (MERC). Mpio Livertador, C. Hornung 84 (MERC). Mpio Sucre, Edo. Mérida, C. Hornung 150 (MERC). Edo. Mérida, Mpio. Libertador, E. Arellano s.n. (MERC). Edo. Mérida, Mpio. Sucre, R. Rico 275 (MERC). Edo. Mérida, Mpio. Sucre, R. Rico 399 (MERC). Edo. Mérida, Laguna de Caparú, A. Rondón 202 (MER). Puya venezuelana L.B.Sm. COLOMBIA: Dpto. del Arauca, Sierra Nevada del Cocuy, Quebrada el Playón, Casa Piedra, A.M. Cleef 10124 (COL). Línea divisoria entre Dpto. Santander del Norte y Cesar, H. García-Barriga 19737 (COL). VENEZUELA: Edo. Mérida, Páramo de Pozo Negro, between San José and Beguilla, J. Steyermark 56285 (F); Edo. Trujillo, L. Aristeguieta 3538 (US). Edo. Mérida, Páramo de Pozo Negro, between San José and Beguilla, J.A. Steyermark 56285 (F). Edo. Mérida, Páramo Gaviria, C. Hornung 204 (MERC). Edo. Mérida, Páramo Gaviria, C. Hornung 205 (MERC). Edo. Mérida, Mpio Sucre, T. Ruiz 44 (MERF). Edo. Mérida, Mpio. Sucre, T. Ruiz 1708 (MERF). Edo. Mérida, Mpio. Libertador, T. Ruiz 8476 (MERF). Puya venusta Phil. ex Baker. CHILE: Gaudichau s.n. (F 741180). Gaudichau s.n. (F 835821). Quilliman 8/9/9 (HDCV). Huentelauquén, poco al sur de la desembocadura del río Choapa, H. González Villalón s.n. 5.xi.1966 (F). Philipii s.n. (F 741266). Prov. Aconcagua, Zapallar, G. Looser 2542 (F). Prov. Aconcagua, Zapallar, G. Looser 2548 (F). Puya westii L.B.Sm. PERÚ: Dpto. La Libertad, Prov. Huamachuco, road to Quiruvilca, 18 km above and west of Huamachuco, P.C. Hutchison 6146 (NY). La libertad, near Huamachuco, J. West 8353 (US). Dpto. La Libertad, Prov. Huamachucho, road to Quiruvilca, P. C. Hutchinson 6146 (F). Tillandsia multicaulis Steud. MÉXICO: Edo. Veracruz, Loma Alta. Mpio. Coatepec, V.E. Luna 1006 (XAL). Edo. Veracruz, Ahuihuixtla, Mpio. Calcahualco, J.L. Martínez 704 (XAL). Edo. Veracruz, Camino Xico a Tonalaco, Mpio Xico, M. Xhazaro 1504 (XAL). Edo. Veracruz, Naolinco, Mpio. Naolinco, F. Ventura 13214 (XAL). Edo. Veracruz, a 2 km de Ahuihutle, camino a tres Aguas (Coscomatepec), Mpio Calcahualco s.n. (XAL). Edo. Veracruz, Alrededores de la represa Xocollolapa, A. Flores-Palacios 941 (XAL). Edo. Veracruz, Jardín Botánico Francisco Javier Clavijero, K. Fabian 346 (XAL). Edo. Veracruz, El Esquilon, Mpio. Jilotepec, R. Ortega 526 (XAL). Veracruz, Tenejapa, S. Avendaño R. 260 (XAL). Edo. Veracruz, J. Martínes G. 187 (XAL). Edo. Veracruz, Etlantepec-Tlacolulan, I. García-Orta 159 (XAL). El Esquilón, Mpio. Jilopetec, M.G. Zola 669 (XAL). Edo. Veracruz, Mpio. Coatepec, V.E. Luna 717 (XAL). Edo. Veracruz, 5 Palos, V.E. Luna M. 840 (XAL). Edo. Veracruz, Cañada río Ayohuxtla, A. Rincón G. 2733 (XAL). Edo. Naranjillos, Mpio. San Andres Tlanehuayocan, Naranjillos, C. Gutierrez 2737 (XAL). Edo. Veracruz, rancho grande, 3 km de Xalapa, J.I. Calzada 1900 (XAL). Edo. Veracruz, along Río Grande, Mun. Las Choapas, M. Nee 29877 (F). Vriesea espinosae (L.B.Sm.) Gilmartin. PERU: Lambayeque, 17 km E of Olmos on road to Pucara, premontane dry forest, A. Gentry 22562 (F). © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110 110 C. T. HORNUNG-LEONI and V. SOSA DATA MATRIX USED FOR CLADISTIC ANALYSIS (CHARACTER NUMBERS CORRESPOND TO THOSE GIVEN IN APPENDIX 2 1 Brocchinia reducta Ananas ananasioides Bromelia pinguin Bromelia chrysantha Aechmea spectabilis Billbergia macrolepis Cottendorfia florida Guzmania monostachia Tillandsia multicaulis Tillandsia flexuosa Vriesea espinosae Navia ignesicola Pitcairnia maidifolia Pitcairnia meridensis Dyckia ferox Puya alpestris Puya berteroniana Puya weddelliana Puya chilensis Puya gilmartiniae Puya boliviensis Puya castellanosii Puya raimondii Puya retrorsa Puya venusta Puya spathacea Puya pygmea Puya coerulea Puya aequatorialis Puya laxa Puya medica Puya floccosa Puya venezuelana Puya aristeguietae Puya trianae Puya goudotiana Puya ferruginea Puya ferreyrae Puya nitida Puya westii Puya nutans Puya santosii Puya cuatrecasasii 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 TABLE 1) 80 87 10––?100210––0011122100011011000110110001001200100001001210100010221000010010111200–031 00––100100102001120110111111001201––––––1––––––100111201200111?01121100111010111212–001 00––13010010100012–1101101111010110010011010011100001001210100010021000010000011201–001 00––13010010100022–20010111210101000100110–0011100101001220100102101400110011001201–001 00––1210011000011211100110111000000100001011010102111201100101012021310001010111201–000 10––110021101001122211001101000000––––––1–––––––02001?002021100101212100111100?0201?000 ?0––1211000–0011122100011011000010100011011011000002?012000000011200001100101010100031 00––1310010––00010–020001100000000––––––1––––––102001$012000100211200001100101$10000010 00––1310010––00110010000000–010001––––––1––––––00001020000001010011010001?000110000?010 00––1211010––001102210011001*0000001000110100001000002101001000011103000100001110002010 00––1111010––001102210010001000001––––––1–––––––0000020010011000?1101?0?1?0?0?1?000?010 010103100110200––1––000?0–––000001––––––1–––––––001–––01–00?1???0?0???0????10??1000?0?1 20––1110000––11011–0100?1000000011––––––1––––––1020012012200011211000001100000000002031 20––1110000––11001––20001000000010––––––1––––––1000011012301010202103100110000200002031 00––?1110012100011–11000110–*100010?10011001000100001001200100021121410010011101010$021 01111211001010011102101010111010110111010001000100000001220020020121201010000011000$121 011*12110012100011–210101–––101011011101001100010200000122$110020021201010000111000?121 011?12010010100??1??10101?1?101011011111000??00?000?0001220?000?1121201110000?11000?121 011112010012000011–21010111210101101111100010101000010012201200201214010100000110001121 01111221001210011112101011121010110111110010010002000101220100020101401010000011000?121 00––12110012100011–11010101010001101110100000001000001112201000221214010100000110002121 0???12010012?0011111101011?21010110111110001011110000?01220*000200?14?1?1?000???0002121 010002010012000$11111020011210111101111100000111000000012201100200210010100000000001121 011112010012000011–21010111$1011010010111000010112101101220100021131$11?10000???000?121 00––112100101001112210110111*00011001001101101010200020122$1101211111111110101010001121 00––1211001010010120100*111110001001100110010111001012012201001201111110100000010001121 00––01110010100111121000001210111100101111000101000101012111100?1121211010000??1000?121 01001211101010011122100111111000110?1001100*01**021012012$01$012211*1111010?00110002121 0111121100112001112210011111000010––––––1––––––100101$012201001211$1$11010000001000?121 00––12110010200221221010111210001011000?1011001100001100230100120121111010000???000?121 01??10110010100011–210000011*000110010111000010102100?012211101201?1211010000010000?121 010012110012100111221001111110001001100110012111000012001101000202211112101001010001121 00––12110011100011–211000012001111––––––1––––––1001001012211100200211110100000110000121 00––11210010000011–11010011$1011110?100111010101000000012201000201210111100000000002121 00––?10100100001110110100112001111––––––1––––––1100001012111100201314110100001010001121 00––12000011000011–2101000121011110?10111000000100000101220100020031$110100001110001121 01***2110012$00$11$210011112*000101100011010000112000100130100022111$011100000$01002121 0???12?1001$0001110210000011000011––––––1––––––100100?01212110120001011010000101000?121 00––12210012000111121000101$000010––––––1––––––1101001012101110201312010000000101002121 00––12?10012000011–110?0????100010?1?00?10110101121011012201000221412110101001010000121 00––?1010010100?11?211000011000111––––––1––––––1101001012201100210314110100000010001121 00––?2010010000011–1101000111011110010111100011110100101221110021131211110000?0?0001121 01?112110010100011–010100012101111001011110001111000010121011002003101111?000101000?121 *, polymorphism (0 and 1); $, polymorphism (1 and 2, or 0 and 2); ?, character unknown; –, not applicable. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 93–110