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Morphological phylogenetics of Puya subgenus Puya (Bromeliaceae)

January 2008
Botanical Journal of the Linnean Society 156(1):93 - 110

January 2008
156(1):93 - 110

DOI:10.1111/j.1095-8339.2007.00740.x

Authors:

Claudia T. Hornung-Leoni

Autonomous University of Hidalgo

Victoria Sosa

Institute of Ecology INECOL

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.

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.

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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].

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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.

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Continuous characters and their logarithmic transformation

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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

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).

The majority of the species belong to subgenus

Puyopsis, and only nine have been included in sub-

genus 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 weddel-

liana Mez, and Puya raimondii Harms. Subsequently,

Puya quillotana W. Weber (1984) and Puya gilmar-

tiniae G.S. Varadarajan & A.R. Flores (1990) were

described. However, the only differentiating character

between P. quillotana and P. chilensis was the pubes-

cence 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.

*Corresponding author. E-mail: victoria.sosa@inecol.edu.mx

Botanical Journal of the Linnean Society, 2008, 156, 93–110. With 4 ﬁgures

Some of these variable characters are found amongst

species of subgenus Puya. For example, the largest

bromeliad in the world, P. raimondii, reaches approxi-

mately 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 well-

developed stem, which may be simple, as in P. rai-

mondii, 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 molecu-

lar 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 Bromeli-

aceae has recognized three subfamilies: Pitcairnio-

ideae, 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 phyloge-

netic 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 sub-

genus Puya until variable molecular characters are

found.

In addition to discrete morphological attributes,

continuous characters were considered in the analy-

ses. There is controversy over the use of these char-

acters 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 continu-

ous 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 rep-

resentative 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 identi-

ﬁed 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, Broc-

chinia was used to root the tree, as it is the most

distantly related genus.

MORPHOLOGICAL DATASET

The data matrix (Appendix 2) consists of 93 charac-

ters: 87 discrete and six continuous characters. Veg-

etative as well as ﬂoral and micromorphological

characters were included. Some of these characters

are shown in Figures 1 and 2. Micromorphological

94 C. T. HORNUNG-LEONI and V. SOSA

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 col-

lected 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 charac-

ters (Farris, 1970). During the optimization of a given

node, if the ranges of the descendant nodes over-

lapped, 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 ANALYSIS

The data matrix was constructed using WINCLADA

(Nixon, 2002). Cladistic analyses under parsimony

criteria were performed using the program TNT

(Goloboff et al., 2003). Separate analyses were per-

formed: the ﬁrst dataset included only the discrete

characters, and the second included the discrete and

continuous characters. Parsimony analyses were per-

formed 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 prob-

ability of 30%. Bremer support (Bremer, 1994) was

calculated only for the combined dataset as imple-

mented in TNT (Goloboff et al., 2003).

RESULTS

CLADISTIC ANALYSES

The analysis with discrete characters retrieved nine

most parsimonious trees (MPTs) [L=505 steps; con-

sistency 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 infor-

mative. In the data matrix with continuous +discrete

characters, 93 characters were parsimony informa-

tive. Autapomorphic characters were scored but not

taken into account in the analyses.

PHYLOGENETIC RELATIONSHIPS

In the discrete character analysis, only a clade that

included two Pitcairnia species was well supported

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.

PHYLOGENY OF PUYA SUBGENUS PUYA 95

96 C. T. HORNUNG-LEONI and V. SOSA

(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 inﬂores-

cence 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 syna-

pomorphic 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 synapo-

morphic 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 speciﬁ-

cally 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 para-

phyletic. 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 res-

urrected. 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 relation-

ships of its species. Such analyses will also clarify the

status of the group formed by northern South Ameri-

can species represented by the P. aristeguietae clade

that is closely related to subgenus Puya.

The topology of the cladograms was better sup-

ported 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 measure-

ments 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

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

PHYLOGENY OF PUYA SUBGENUS PUYA 97

Table 1. Characters and character states for the cladistic analysis of subgenus Puya

1. Plant length

2. Leaf length

3. Inﬂorescence length

4. Floral bract length

5. Sepal length

6. Petal length

7. 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

8. Stem: 0, acaulescent; 1, caulescent

9. Stem type: 0, erect; 1, prostrate

10. Stem branching: 0, simple; 1, branched

11. Reproduction: 0, monocarpic; 1, polycarpic

12. 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

13. Leaf position in the rosette during blooming: 0, erect; 1, recurved-reﬂexed; 2, ascendent

14. Leaf margin: 0, parallel; 1, convergent

15. Leaf blade apex: 0, acute or attenuate; 1, rounded; 2, mucronate or acuminate

16. Leaf capacity of leaves to retain water: 0, absent; 1, present

17. Leaf margin: 0, entire; 1, serrate

18. Leaf margin: thorn orientation: 0, one way, antrorse; 1, one way, retrorse; 2, with both orientations, antrorse

and retrorse

19. Leaf margin: thorn colour: 0, black-brownish; 1, brown-reddish or greenish; 2, yellow to red

20. Leaf nervation: 0, without a prominent median nerve; 1, with a prominent median nerve

21. Leaf petiole: 0, absent; 1, present

22. Leaf pubescence of adaxial surface: 0, glabrous; 1, with scales; 2, with hairs (pubescent)

23. Leaf pubescence of abaxial surface: 0, glabrous; 1, with scales; 2, with hairs (pubescent)

24. 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

25. Leaf adaxial surface: percentage of pubescence: 0, 25%; 1, 50–75%; 2, 100%

26. Leaf abaxial surface: percentage of pubescence: 0, 25%; 1, 50–75%; 2, 100%

27. Inﬂorescence: peduncle: 0, included in rosette; 1, emerging rosette; 2, equalling rosette

28. Inﬂorescence: 0, erect; 1, pendule or nutant

29. Inﬂorescence: peduncle diameter: 0, less than 2 cm; 1, 2–10 cm; 2, more than 10 cm

30. Inﬂorescence reddish coloration: 0, absent; 1, present

31. Inﬂorescence: peduncle covered with bracts: 0, totally; 1, partially

32. Inﬂorescence: peduncle bracts shape: 0, oblong-elliptic; 1, ovate

33. Inﬂorescence: bract margin: 0, entire; 1, serrate or serrulate

34. Inﬂorescence: bract pubescence: 0, glabrous; 1, with scales; 2, with hairs

35. Inﬂorescence: branching: 0, simple; 1, branched

36. Inﬂorescence position: 0, apical; 1, lateral

37. Inﬂorescence: consistency of the rachis of inﬂorescence: 0, nonﬂeshy; 1, ﬂeshy

38. Inﬂorescence: strobiliform: 0, absent; 1, present

39. Inﬂorescence: ﬂower disposition type: 0, spikes; 1, racemes

40. Inﬂorescence: ﬂower density in the entire inﬂorescence: 0, lax; 1, dense

41. Inﬂorescence: branch arrangement: 0, polystichous; 1, distichous

42. Inﬂorescence: branch shape: 0, globose; 1, oblong-elliptic

43. Inﬂorescence: branch position: 0, alternate; 1, verticillate

44. Inﬂorescence: sterile apex of branches: 0, absent; 1, present

45. Inﬂorescence: density of ﬂowers along branches: 0, lax; 1, dense

46. Inﬂorescence: shape: 0, triangular; 1, oblong

47. Inﬂorescence: blooming pattern: 0, ﬂowers opening sequentially from inﬂorescence branch base to apex; 1,

ﬂowers opening in any position

48. Inﬂorescence: bract/branch length: 0, bracts smaller or equalling branches; 1, bracts larger

49. 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

98 C. T. HORNUNG-LEONI and V. SOSA

exception of a small terminal clade formed by P. cas-

tellanosii 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 compari-

son 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 sug-

gested 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

Table 1. Continued

50. Inﬂorescence: bract shape: 0, oblong-elliptic; 1, ovate

51. Inﬂorescence: apical zone of the inﬂorescence bracts apex: 0, acute-attenuate; 1, rounded; 2,

mucronate-apiculate

52. Inﬂorescence: margin of bract: 0, entire; 1, serrate or serrulate

53. Inﬂorescence: blade of bracts: 0, absent; 1, present

54. Inﬂorescence rachis pubescent: 0, absent; 1, present

55. Inﬂorescence: ferrugineous pubescence: 0, absent; 1, present

56. Inﬂorescence: bract apex: 0, acute-attenuate; 2, rounded or obtuse; 2, mucronate or apiculate

57. Floral bract: margin: 0, entire; 1, serrate or serrulate

58. Floral bract: carinate: 0, absent; 1, present

59. Floral bract shape: 0, oblong-elliptic; 1, ovate

60. Floral bract colour (fresh material): 0, greenish; 1, brownish; 2, red–orange

61. Floral sterile bracts among ﬂowers: 0, absent; 1, present

62. Flower disposition: 0, distichous; 1, polystichous

63. Flower phyllotaxis: 0, opposite; 1, alternate; 2, verticillate

64. Flower: pedicel: 0, absent; 1, short (<2 mm); 2, medium (2–6 cm); 3, large (>6 cm)

65. Flower: length with regard to bracts: 0, larger; 1, smaller; 2, equal size

66. Flower: bract pubescence: 0, absent; 1, present

67. Flower: sepal length with regard to bracts: 0, larger; 1, smaller; 2, equal size

68. Flower: sepal symmetry: 0, symmetrical; 1, strongly asymmetrical

69. Flower: sepals carinate: 0, absent; 1, present

70. Flower: sepal union: 0, free; 1, sepals connate at least at half; 2, connate only at the base

71. Flower: sepal apex: 0, acute-attenuate; 1, rounded-obtuse; 2, mucronate-acuminate

72. Flower: sepal length with regard to petal: 0, 1/3 smaller; 1, 1/2 smaller or less; 2, equal size

73. Flower: sepal colour: 0, yellowish; 1, reddish or coloured; 2, greenish-white–cream; 3, brownish

74. Flower: sepal pubescence: 0, absent; 1, present

75. Flower: petal colour: 0, white–cream; 1, blue–violet; 2, dark green–blue; 3, red–rose; 4, yellow-greenish

76. Flower: petal shape: 0, oblong-elliptic; 1, obovate

77. Flower: petals spiralled and twisted together after anthesis: 0, absent; 1, present

78. Flower: petal apex: 0, acute-attenuate; 1, rounded-obtuse; 2, mucronate-acuminate

79. Flower: petal symmetry: 0, asymmetric; 1, symmetric

80. Flower: petaloid appendices at base of petals: 0, absent; 1, present

81. Flower: relative size of stamen with respect to the ﬂowers: 0, inserted; 1, exerted

82. Flower: stamens adnate to petals: 0, absent; 1, present

83. Flower: stamens in a column: 0, absent; 1, present

84. Flower: anthers: 0, basiﬁxed; 1, dorsiﬁxed

85. Flower: stamen length with regard to style: 0, larger or equal size; 1, smaller

86. Flower: anther shape: 0, sagittate; 1, oblong-elliptic

87. Flower: ovary: 0, superior; 1, semi-inferior; 2, inferior

88. Flower: style size with regard to ovary: 0, larger; 1, smaller or equal

89. Fruit: type: 0, capsule; 1, berry; 2, drupe; 3, multiple fruit

90. Fruit: dehiscence of capsules: 0, septicidal; 1, loculicidal; 2, both

91. Fruit: petals persistent at apex of fruit: 0, absent; 1, present

92. Seed: 0, naked; 1, plumose; 2, winged; 3, appendiculate

93. Roots: wide radicular system: 0, absent; 1, present

PHYLOGENY OF PUYA SUBGENUS PUYA 99

Table 2. Continuous characters and their logarithmic transformation

Species

012 345

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 99–100 2.00 159–161 2.21 14.5–15.0 1.19–1.20 1.5–2.0 0.40–0.48 0.65–0.7 0.22–0.23 1.3–1.7 0.36–0.43

Ananas ananasioides 99–101 2.00 160.0 2.21 14.5–15.0 1.19–1.20 1.5–2.0 0.40–0.48 0.65–0.7 0.22–0.23 1.3–1.7 0.36–0.43

Bromelia penguin 100–200 2.00–2.30 30.0–200.0 1.49–2.30 30.0–60.0 1.49–1.79 1.6–3.0 0.41–0.60 1.2–3.0 0.34–0.60 1.5–3.0 0.40–0.60

Broemlia chrysantha 70–150 1.85–2.18 80.0–150.0 1.91–2.18 79.0–81.0 1.90–1.91 1.0–1.5 0.30–0.40 0.9–1.7 0.28–0.43 0.9–1.5 0.28–0.40

Aechmea spectabilis 99–101 2.00–2.01 100.0–140.0 2.00–2.15 60.0–100.0 1.79–2.00 0.1–0.2 0.04–0.04 0.9–2.0 0.28–0.48 2.4–2.5 0.53–0.54

Billbergia macrolepis 98–102 2.00–2.01 119–120 2.08 39.5–40.0 161.00 3.45–3.5 0.65 0.99–1.0 0.30 4.2–4.3 0.72

Cottendorﬁa ﬂorida 200–400 2.30–2.60 99–100 2.00 100.0–300.0 2.00–2.48 0.19–0.2 0.08 0.28–0.3 0.11 0.6–1.0 0.20–0.30

Guzmania monostachia 20–40 1.32–1.61 16.0–24.0 1.23–1.40 8.0–15.0 0.95–1.20 2.3–2.9 0.52–0.59 1.75–1.80 0.44–0.45 3.0–3.3 0.60–0.63

Tillandsia multicaulis 25–40 1.41–1.61 30.0–40.0 1.49–1.61 14.0–14.5 1.18–1.19 4.9–5.0 0.77–0.78 3.5–3.6 0.65–0.66 6.9–7.0 0.90

Tillandsia ﬂexuosa 20–150 1.32–2.18 20.0–50.0 1.32–1.71 50–61 1.78–1.79 2.0–3.0 0.48–0.60 2.0–3.0 0.48–0.60 3.9–4 0.69–0.70

Vriesea espinosae 15–16 1.20–1.23 ? ? 6.9–7 0.90 2.4–2.5 0.53–0.54 1.19–1.2 0.34 2.9–3.0 0.59–0.60

Navia ignesicola 16.5–17 1.25–1.26 39–41 1.60–1.62 4.9–5.0 0.77–0.78 2.6–2.7 0.56–0.57 4.4–4.5 0.73–0.74 ? ?

Pitcairnia maidifolia 128–131 2.11–2.12 119–120 2.08 44–45 1.65–1.66 0.39–0.40 0.14–0.15 2.6–3.0 0.56–0.60 4.9–5.0 0.77–0.78

Pitcairnia meridensis 50–60 1.71–1.79 70.0–130.0 1.85–2.12 14.9–15 1.20 0.7–1.1 0.23–0.32 3.1–3.2 0.61–0.62 6.35–6.40 0.87

Dyckia ferox 58–62 1.77–1.79 20–60 1.32–1.79 ? ? 0.2–0.3 0.08–0.11 0.5–0.9 0.18–0.28 1.2–1.4 0.34–0.38

Puya alpestris 120–150 2.08–2.18 60–70 1.79–1.85 99–100 2.00 2.6–3.3 0.56–0.63 2.2–2.9 0.51–0.59 4.8–5.0 0.76–0.78

Puya berteroniana 495–505 2.70 99–100 2.00 99–100 2.00 4.4–4.8 0.73–0.76 2.2–2.3 0.51–0.52 5.0–5.6 0.78–0.82

Puya weddelliana ? ? ? ? ? ? 1.49–1.5 0.40 1.79–1.80 0.45 3.45–3.50 0.65

Puya chilensis 499–500 2.70 99–100 2.00 99–100 2.00 3.4–5.0 0.64–0.78 3.5–5.8 0.65–0.83 4.9–5.0 0.77–0.78

Puya gilmartiniae 149–151 2.18 10.0–20.0 1.04–1.32 50.0–60.0 1.71–1.79 1.5–2.5 0.40–0.54 1.7–2.4 0.43–0.53 3.0–4.8 0.60–0.76

Puya boliviensis 199–200 2.30 100.0 2.00 99–100 2.00 2.1–3.3 0.49–0.63 3.19–3.20 0.62 4.9–5.0 0.77–0.78

Puya castellanosii 100–300 2.00–2.48 39–40 1.60–1.61 39–40 1.60–1.61 3.2–3.5 0.62–0.65 2.9–3.0 0.59–0.60 3.9–4.0 0.69–0.70

Puya raimondii 950–1200 2.98–3.08 123–125 2.09–2.10 430–500 2.63–2.70 3.1–6.0 0.61–0.85 4.0–4.1 0.70–0.71 7.9–8.0 0.95

Puya retrorsa 200–300 2.30–2.48 40.0–60.0 1.61–1.79 40.0–80.0 1.61–1.69 3.0–3.5 0.60–0.65 1.99–2.0 0.48 3.7–4.0 0.67–0.70

Puya venusta 90–100 1.96–2.00 30.0–35.0 1.49–1.56 30.0–35.0 1.49–1.56 0.39–0.4 0.14–0.15 1.5–2.0 0.40–0.48 3.4–3.5 0.64–0.65

Puya spathacea 99–100 2.00 60.0–100.0 1.79–2.00 40.0–60.0 1.61–1.79 1.3–2.2 0.36–0.51 1.5–2.2 0.40–0.51 2.5–3.3 0.54–0.63

Puya pygmea 20–30 1.32 15.0–22.0 1.20–1.36 3.0–5.0 0.60–0.78 2.9–3.0 0.59–0.60 1.5–1.8 0.40–0.45 1.5–2.0 0.40–0.48

Puya coerulea 199–200 2.30 59–61 1.78–1.79 98–101 2.00–2.01 1.69–1.7 0.43 2.39–2.40 0.53 3.7–5.0 0.67–0.78

Puya aequatorialis 199–200 2.30 99–100 2.00 98–101 2.00–2.01 1.79–1.8 0.45 2.0–2.3 0.48–0.52 2.9–3.0 0.59–0.60

Puya laxa 79–81 1.90–1.91 27–28 1.45–1.46 70.0–80.0 1.85–1.91 1.3–2.0 0.36–0.48 1.69–1.70 0.43 2.9–3.0 0.59–0.60

Puya medica 20–37 1.32–1.58 20–20.5 1.32–1.33 10.0–20.0 1.04–1.32 1.5–1.6 0.40–0.41 1.8–2.0 0.45–0.48 3.4–4.0 0.64–0.70

Puya ﬂoccosa 50–200 1.71–2.30 100.0–110.0 2.00–2.05 30.0–100.0 1.49–2.00 1.3–1.7 0.36–0.43 2.5–3.3 0.54–0.63 3.9–4.0 0.69–0.70

Puya venezuelana 59–60 1.78–1.79 25.0–34.0 1.41–1.54 18.0–22.0 1.28–1.36 4.45–4.5 0.74 1.8–2.0 0.45–0.48 3.0–3.5 0.60–0.65

Puya aristeguietae 299–300 2.48 99–100 2.00 100.0–110.0 2.00–2.05 1.6–1.8 0.41–0.45 2.5–3.3 0.54–0.63 5.0–6.0 0.78–0.85

Puya trianae 40–200 1.61–2.30 24.0–28.0 1.40–1.46 15.0–30.0 1.20–1.49 5.0–6.0 0.78–0.85 2.0–2.5 0.48–0.54 3.10–3.20 0.61–0.62

Puya goudotiana 499–500 2.70 100.0–170.0 2.00–2.23 100.0–200.0 2.00–2.30 2.0–3.7 0.48–0.67 2.0–3.0 0.48–0.60 5.0–6.0 0.78–0.85

Puya ferruginea 250–400 2.40–2.60 99–100 2.00 199–200 2.30 4.45–4.5 0.74 1.2–4.5 0.34–0.74 8.0–14.0 0.95–1.18

Puya ferreyrae 150–200 2.18–2.30 55.0–100.0 1.75–2.00 60.0–75.0 1.79–1.88 5.0–7.0 0.78–0.90 3.2–3.7 0.62–0.67 3.9–4.0 0.69–0.70

Puya nitida 170–200 2.23–2.30 35.0–60.0 1.56–1.79 58.0–62.0 1.77–1.80 3.4–3.8 0.64–0.68 2.9–3.0 0.59–0.60 6.9–7.0 0.90

Puya westii 249–250 2.40 99–100 2.00 49–50 1.70–1.71 0.8–1.0 0.26–0.30 1.99–2.0 0.48 3.9–4.0 0.69–0.70

Puya nutans 55–70 1.75–1.85 15.0–23.5 1.20–1.39 29–30 1.48–1.49 1.99–2.0 0.48 1.79–1.80 0.45 3.9–4.0 0.69–0.70

Puya santosii 80–200 1.91–2.30 31.0–37.0 1.51–1.58 32.0–50.0 1.52–1.71 3.2–3.3 0.62–0.63 1.3–2.0 0.36–0.48 3.0–3.6 0.60–0.66

Puya cuatrecasasii 99–100 2.00 25–27 1.43 ? ? 2.59–2.6 0.56 2.35–2.4 0.53 4.90–5.0 0.77–0.78

100 C. T. HORNUNG-LEONI and V. SOSA

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 medium-

sized, 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 Lati-

noamericana 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

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].

PHYLOGENY OF PUYA SUBGENUS PUYA 101

due to J. Betancur, J. Manzanares, J. R. Grant, M.

Dillon, and B. Holst for their advice, and to G. Mon-

tenegro, 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.

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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). COLOM-

BIA: 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).

PHYLOGENY OF PUYA SUBGENUS PUYA 103

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.DoRon-

donia, Mpio. De Ariquemes, Mineracao Mibrasa,

Setor Alto Candeias, km 128, Sudoeste de Ariquemes,

L.O.A.Teixeira 611(NY).V

ENEZUELA: Edo. Amazonas,

Cerro Sipapo (Paráque), B. Maguire 28053 (NY). Bajo

el Río Guaramacal, R. Liesner 10610 (VEN).

Brocchinia acuminata L.B.Sm. VENEZUELA: Edo. Ter-

ritorio 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. Castillo-

Campos 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. VEN-

EZUELA: Territorio Federal Amazonas, Dpto. Atures:

forested areas and igneous outcrops along Río Coro-

moto, at Tobogán de la Selva, J. Steyermark 122478

(US).

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,

104 C. T. HORNUNG-LEONI and V. SOSA

O. Valverde 515 (F). Prov. San José, F. Montgomery

G. 5052 (F). GUIANAS: Region Cuyuni-Mazaruni, Mt.

Holitipu, below peak, T. McDowell 3027 (F). HONDU-

RAS: 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.Will-

iams 42265 (F). Dpto. Morazán, Region of Agua Ama-

rilla, 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

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 Brace-

lin 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. Wed-

ermann 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-

PHYLOGENY OF PUYA SUBGENUS PUYA 105

enda Rinconada La Cesda, Maipu, Qda La Plata, F.

Schlegel 1664 (SGO). Prov. Santiago, Región Metro-

politana, Reserva Forestal Río Clarillo, J. Yañez s.n.

3.viii.1993 (SGO). Prov. Valparaiso, four km south-

west 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). Ran-

cagua, 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: Antofa-

gasta 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. Vara-

darajan 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): Gaud-

ichaud, 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 Col-

chagua, 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). Que-

brada 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. Cuatreca-

sas 27573 (COL). Cordillera Central, vertiente occi-

dental, J. Cuatrecasas 18962 (COL). Dpto. Cauca,

Macizo Colombiano, Páramo de las Papas, J. Idrobo

4062 (COL). Dpto. del Cauca, Cordillera Central, ver-

tiente occidental, Cabeceras del Río Palo. J. Cuatre-

casas 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. Cajama-

rca, 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,SC634, 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

106 C. T. HORNUNG-LEONI and V. SOSA

(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, Neque-

jahuira, 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. Huaylla-

bamba, 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 (Collambay-

Sinsicap), A. Sagástegui 15638 (F). Lima, Prov. Hua-

rochiri, 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,

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. Cundinama-

rca, 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. Cuatreca-

sas 13547 (F). Dpto. Norte de Santander, Cordillera

Oriental, J. Cuatrecasas 10039 (F). Dpto. Cundi-

namarca, 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. Amazo-

nas, Páramo de Guasca, A. Fernández P. 5760

PHYLOGENY OF PUYA SUBGENUS PUYA 107

(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. Cuatreca-

sas 13539 (F). Macizo de Bogotá, Cerro entre que-

brada de Las Delicias y la de Las Ninfas, J.

Cuatrecasas 5629 (F). Dpto. Cundinamarca, Cood-

illera 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 Guata-

vita, J. Betancur 2713 (COL, NY). Dpto. Cundinama-

rca, 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 characteris-

tics 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). Weber-

bauer 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, Huishcash-

pampa, E. Cerrate 2072 (USM). Prov. Huaylas, Dpto.

Ancash, A. Cano 6405 (USM). 1/2 km SE of Haci-

enda Santa Rosa de Achaco, s.n. (USM 159980).

Dpto. Huancavelica, carretera Castrovirreyna-

Ayacucho, 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. Chimbo-

razo, between Cajabamba and Pallatanga, G.S. Vara-

darajan 1440 (US). Prov. Pichincha, Paramo de

Huamani, G.S. Varadarajan 1420 (US). Tunguragua,

A.J. Gilmartin 1103 (US).

Puya santosii Cuatrec. COLOMBIA: Dpto. Cundinama-

rca, Cordillera Oriental, Páramo de Cruz Verde,

J. Cuatrecasas 9518 (F). Dpto. de Cundinamarca,

Macizo de Bogotá, Páramo de Usaquén, J. Cuatreca-

sas 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. Cundi-

namarca, Macizo de Bogotá, Páramo de Uraquén, J.

Cuatrecasas 9441 (COL). Meta: Páramo de Sumapáz,

Hoya El Nevado, A. Cleef 1503 (COL). Dpto. Cundi-

namarca, 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. Cundinama-

rca, Cordillera Oriental, J. Cuatrecasas 9518 (F).

Puya spathacea (Griseb.) Mez. ARGENTINA:Prov.De

Cordoba, Sierra Grande, A.P. Rodrigo 458 (NY).

108 C. T. HORNUNG-LEONI and V. SOSA

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 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. Cundinama-

rca, 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, Ser-

ranía Peña Negra, A.M. Cleef 7430 (COL). VENEZU-

ELA: Edo. Mérida, Mpio. Campo Elías, T.Ruiz 8000

(MERF).

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. Acon-

cagua, Zapallar, G. Looser 2542 (F). Prov. Aconcagua,

Zapallar, G. Looser 2548 (F).

Puya weddelliana Mez. BOLIVIA: Dpto. Chuquisaca,

D.S. Correll 644 (US). Tarija, Weddell 4001 (US

photo).

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 ﬂexuosa Sw. COLOMBIA: Vaupes, Río Kudu-

yarí, Yapobodá, sandstone savannah near headwa-

ters, 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).

Tillandsia multicaulis Steud. MÉXICO: Edo. Ver-

acruz, Loma Alta. Mpio. Coatepec, V.E. Luna 1006

(XAL). Edo. Veracruz, Ahuihuixtla, Mpio. Calcahua-

lco, 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 Cal-

cahualco 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). Ver-

acruz, 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: Lam-

bayeque, 17 km E of Olmos on road to Pucara, pre-

montane dry forest, A. Gentry 22562 (F).

PHYLOGENY OF PUYA SUBGENUS PUYA 109

APPENDIX 2

DATA MATRIX USED FOR CLADISTIC ANALYSIS (CHARACTER NUMBERS CORRESPOND TO THOSE GIVEN IN TABLE 1)

1 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 87

Brocchinia

reducta

10––?100210––0011122100011011000110110001001200100001001210100010221000010010111200–031

Ananas

ananasioides

00––100100102001120110111111001201––––––1––––––100111201200111?01121100111010111212–001

Bromelia

pinguin

00––13010010100012–1101101111010110010011010011100001001210100010021000010000011201–001

Bromelia

chrysantha

00––13010010100022–20010111210101000100110–0011100101001220100102101400110011001201–001

Aechmea

spectabilis

00––1210011000011211100110111000000100001011010102111201100101012021310001010111201–000

Billbergia

macrolepis

10––110021101001122211001101000000––––––1–––––––02001?002021100101212100111100?0201?000

Cottendorﬁa

ﬂorida

?0––1211000–0011122100011011000010100011011011000002?012000000011200001100101010100031

Guzmania

monostachia

00––1310010––00010–020001100000000––––––1––––––102001$012000100211200001100101$10000010

Tillandsia

multicaulis

00––1310010––00110010000000–010001––––––1––––––00001020000001010011010001?000110000?010

Tillandsia

ﬂexuosa

00––1211010––001102210011001*0000001000110100001000002101001000011103000100001110002010

Vriesea

espinosae

00––1111010––001102210010001000001––––––1–––––––0000020010011000?1101?0?1?0?0?1?000?010

Navia

ignesicola

010103100110200––1––000?0–––000001––––––1–––––––001–––01–00?1???0?0???0????10??1000?0?1

Pitcairnia

maidifolia

20––1110000––11011–0100?1000000011––––––1––––––1020012012200011211000001100000000002031

Pitcairnia

meridensis

20––1110000––11001––20001000000010––––––1––––––1000011012301010202103100110000200002031

Dyckia ferox 00––?1110012100011–11000110–*100010?10011001000100001001200100021121410010011101010$021

Puya alpestris 01111211001010011102101010111010110111010001000100000001220020020121201010000011000$121

Puya

berteroniana

011*12110012100011–210101–––101011011101001100010200000122$110020021201010000111000?121

Puya

weddelliana

011?12010010100??1??10101?1?101011011111000??00?000?0001220?000?1121201110000?11000?121

Puya chilensis 011112010012000011–21010111210101101111100010101000010012201200201214010100000110001121

Puya

gilmartiniae

01111221001210011112101011121010110111110010010002000101220100020101401010000011000?121

Puya

boliviensis

00––12110012100011–11010101010001101110100000001000001112201000221214010100000110002121

Puya

castellanosii

0???12010012?0011111101011?21010110111110001011110000?01220*000200?14?1?1?000???0002121

Puya

raimondii

010002010012000$11111020011210111101111100000111000000012201100200210010100000000001121

Puya retrorsa 011112010012000011–21010111$1011010010111000010112101101220100021131$11?10000???000?121

Puya venusta 00––112100101001112210110111*00011001001101101010200020122$1101211111111110101010001121

Puya

spathacea

00––1211001010010120100*111110001001100110010111001012012201001201111110100000010001121

Puya pygmea 00––01110010100111121000001210111100101111000101000101012111100?1121211010000??1000?121

Puya coerulea 01001211101010011122100111111000110?1001100*01**021012012$01$012211*1111010?00110002121

Puya

aequatorialis

0111121100112001112210011111000010––––––1––––––100101$012201001211$1$11010000001000?121

Puya laxa 00––12110010200221221010111210001011000?1011001100001100230100120121111010000???000?121

Puya medica 01??10110010100011–210000011*000110010111000010102100?012211101201?1211010000010000?121

Puya ﬂoccosa 010012110012100111221001111110001001100110012111000012001101000202211112101001010001121

Puya

venezuelana

00––12110011100011–211000012001111––––––1––––––1001001012211100200211110100000110000121

Puya

aristeguietae

00––11210010000011–11010011$1011110?100111010101000000012201000201210111100000000002121

Puya trianae 00––?10100100001110110100112001111––––––1––––––1100001012111100201314110100001010001121

Puya

goudotiana

00––12000011000011–2101000121011110?10111000000100000101220100020031$110100001110001121

Puya

ferruginea

01***2110012$00$11$210011112*000101100011010000112000100130100022111$011100000$01002121

Puya ferreyrae 0???12?1001$0001110210000011000011––––––1––––––100100?01212110120001011010000101000?121

Puya nitida 00––12210012000111121000101$000010––––––1––––––1101001012101110201312010000000101002121

Puya westii 00––12?10012000011–110?0????100010?1?00?10110101121011012201000221412110101001010000121

Puya nutans 00––?1010010100?11?211000011000111––––––1––––––1101001012201100210314110100000010001121

Puya santosii 00––?2010010000011–1101000111011110010111100011110100101221110021131211110000?0?0001121

Puya

cuatrecasasii

01?112110010100011–010100012101111001011110001111000010121011002003101111?000101000?121

*, polymorphism (0 and 1); $, polymorphism (1 and 2, or 0 and 2); ?, character unknown; –, not applicable.

110 C. T. HORNUNG-LEONI and V. SOSA

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Full-text available

Jul 2022

The Andean plant endemic Puya is a striking example of recent and rapid diversifica-tion from central Chile to the northern Andes, tracking mountain uplift. This study generated 12 complete plastomes representing nine Puya species and compared them to five published plastomes for their features, genomic evolution, and phylogeny. The total size of the Puya plastomes ranged from 159,542 to 159,839 bp with 37.3%– 37.4% GC content. The Puya plastomes were highly conserved in organization and structure with a typical quadripartite genome structure. Each of the 17 consensus plastomes harbored 133 genes, including 87 protein-coding genes, 38 tRNA (transfer RNA) genes, and eight rRNA (ribosomal RNA) genes; we found 69– 78 tandem repeats, 45– 60 SSRs (simple sequence repeats), and 8– 22 repeat structures among 13 spe-cies. Four protein-coding genes were identified under positive site-specific selection in Puya. The complete plastomes and hypervariable regions collectively provided pro-nounced species discrimination in Puya and a practical tool for future phylogenetic studies. The reconstructed phylogeny and estimated divergence time for the lineage suggest that the diversification of Puya is related to Andean orogeny and Pleistocene climatic oscillations. This study provides plastome resources for species delimitation and novel phylogenetic and biogeographic studies.

Tesis Doctoral: Filogenia y evolución de la subtribu Melinidinae (Paniceae: Panicoideae: Poaceae)

Thesis

Full-text available

Nov 2010

Diego L. Salariato

La subtribu Melinidinae (Poaceae: Panicoideae: Paniceae) incluye 13 géneros y aproximadamente 190 especies, las cuales se hallan caracterizadas por presentar la vía fotosintética C4 PCK y una anatomía foliar distintiva. El objetivo de este trabajo es estudiar las relaciones filogenéticas y la historia evolutiva de este grupo. Para esto se propone analizar a la subtribu desde tres diferentes niveles taxonómicos. En una primera parte se estudia la relación de Melinidinae con los demás integrantes de la tribu Paniceae, poniendo a prueba su monofilia, resolviendo la relación entre la subtribu y su grupo hermano y reconstruyendo la vía fotosintética ancestral. En una segunda parte se estudian las relaciones filogenéticas entre los distintos géneros incluidos en Melinidinae y se reconstruye la tendencia evolutiva que presenta la subtribu en cuanto al grado de homogenización de sus inflorescencias. Finalmente, se analiza en la tercera parte del trabajo las relaciones existentes entre las especies de Urochloa, género con mayor número de especies dentro de la subtribu, con el objetivo de definir grupos infragenéricos naturales y caracteres morfológicos de valor taxonómico. En cada sección se utilizan métodos de análisis y caracteres adecuados para el nivel taxonómico que se estudia. Los resultados obtenidos muestran que la subtribu Melinidinae resulta monofilética cuando Urochloa venosa es excluida de ésta, mientras que la asignación del grupo y del tipo fotosintético ancestral son ambiguas. Los géneros Urochloa y Eriochloa resultan parafileticos y/o polifiléticos, mientras que géneros como Megathyrsus, Melinis y Moorochloa son monofiléticos. La homogenización de las inflorescencias en la subtribu parece ser la tendencia evolutiva favorecida por sobre el proceso inverso. Finalmente, el género Urochloa es, en su actual circunscripción, un grupo artificial carente de sinapomorfías morfológicas. No obstante, los análisis recuperan dentro de Urochloa varios grupos infragénericos definidos por diferentes caracteres morfológicos. Sobre la base de estos resultados se establece para las especies Urochloa acuminata, U. decidua y U. megastachya el nuevo género Rupichloa.

Re-establishment of Aechmea laxiflora (Bromeliaceae, Bromelioideae) based on morphological and phylogenetic evidence, with notes on its geographical distribution

Article

Full-text available

Mar 2024

Background: Previous phylogenetic analyses suggested that Aechmea bracteata is not a monophyletic species, being each variety an individual lineage within the Aechmea bracteata complex. Hypothesis: A phylogenetic analysis based on molecular data and a morphological analysis will provide evidence to support the taxonomic recognition of A. bracteata var. pacifica as a distinct species. Studies species: A. bracteata var. bracteata, A. bracteata var. pacifica and related species. Study site and dates: Mexico to Northern South America; 17 populations from Southeastern and Western Mexico. Analyses were performed between 2021 and 2023. Methods: A linear morphometric analysis was conducted with 85 herbarium specimens testing 28 quantitative and three qualitative variables. Potential diagnostic traits were reviewed in living specimens. A phylogenetic analysis was performed with two nDNA markers (ETS and g3pdh) and one cpDNA marker (trnL-F). A total of 373 records were projected into biogeographical provinces of the Neotropics. Results: The morphometric analysis allowed to separate each variety into a group, also, nine of the evaluated traits resulted statistically significant through a univariate analysis. Five additional diagnostic traits from the inflorescence and leaves were recognized. According to nDNA both taxa present reciprocal monophyly, however, cpDNA groups A. bracteata var. pacifica with Central American species, showcasing a potential hybrid origin. Geographically, the Sierra Madre del Sur acts as a barrier between both taxa. Conclusions: Due to the gathered evidence is possible to recognize A. bracteata var. pacifica as an endemic species to Mexico, reestablishing the basionym A. laxiflora.

Phylogenetic analysis of a new morphological dataset elucidates the evolutionary history of Crocodylia and resolves the long-standing gharial problem

Article

Full-text available

Sep 2021

First appearing in the latest Cretaceous, Crocodylia is a clade of semi-aquatic, predatory reptiles, defined by the last common ancestor of extant alligators, caimans, crocodiles, and gharials. Despite large strides in resolving crocodylian interrelationships over the last three decades, several outstanding problems persist in crocodylian systematics. Most notably, there has been persistent discordance between morphological and molecular datasets surrounding the affinities of the extant gharials, Gavialis gangeticus and Tomistoma schlegelii . Whereas molecular data consistently support a sister taxon relationship, in which they are more closely related to crocodylids than to alligatorids, morphological data indicate that Gavialis is the sister taxon to all other extant crocodylians. Here we present a new morphological dataset for Crocodylia based on a critical reappraisal of published crocodylian character data matrices and extensive firsthand observations of a global sample of crocodylians. This comprises the most taxonomically comprehensive crocodylian dataset to date (144 OTUs scored for 330 characters) and includes a new, illustrated character list with modifications to the construction and scoring of characters, and 46 novel characters. Under a maximum parsimony framework, our analyses robustly recover Gavialis as more closely related to Tomistoma than to other extant crocodylians for the first time based on morphology alone. This result is recovered regardless of the weighting strategy and treatment of quantitative characters. However, analyses using continuous characters and extended implied weighting (with high k -values) produced the most resolved, well-supported, and stratigraphically congruent topologies overall. Resolution of the gharial problem reveals that: (1) several gavialoids lack plesiomorphic features that formerly drew them towards the stem of Crocodylia; and (2) more widespread similarities occur between species traditionally divided into tomistomines and gavialoids, with these interpreted here as homology rather than homoplasy. There remains significant temporal incongruence regarding the inferred divergence timing of the extant gharials, indicating that several putative gavialids (‘thoracosaurs’) are incorrectly placed and require future re-appraisal. New alligatoroid interrelationships include: (1) support for a North American origin of Caimaninae in the latest Cretaceous; (2) the recovery of the early Paleogene South American taxon Eocaiman as a ‘basal’ alligatoroid; and (3) the paraphyly of the Cenozoic European taxon Diplocynodon . Among crocodyloids, notable results include modifications to the taxonomic content of Mekosuchinae, including biogeographic affinities of this clade with latest Cretaceous–early Paleogene Asian crocodyloids. In light of our new results, we provide a comprehensive review of the evolutionary and biogeographic history of Crocodylia, which included multiple instances of transoceanic and continental dispersal.

Nuevos registros de Puya (Bromeliaceae) en el departamento de Lambayeque, Perú

Article

Full-text available

May 2021

Este estudio proporciona una lista comentada de las especies de Puya (Bromeliaceae) registradas en el departamento de Lambayeque, Perú. Se reportan un total de cinco especies, de las cuales cuatro se registran por primera vez y una de ellas se confirma como endemismo de los Andes del Departamento. Las ecorregiones donde se encuentran corresponden al Bosque Seco estacional (BSe), la transición Matorral Desértico-Bosque Seco (MD-BS) y Jalca (JA). Se presenta una clave para las especies reportadas y se discuten las razones de los endemismos, amenazas y sus criterios de categorización.

Ecosystem services show variable responses to future climate conditions in the Colombian páramos

Article

Full-text available

May 2021

Background The páramos, the high-elevation ecosystems of the northern Andes, are well-known for their high species richness and provide a variety of ecosystem services to local subsistence-based communities and regional urbanizations. Climate change is expected to negatively affect the provision of these services, but the level of this impact is still unclear. Here we assess future climate change impact on the ecosystem services provided by the critically important páramos of the department of Boyacá in Colombia, of which over 25% of its territory is páramo. Methods We first performed an extensive literature review to identify useful species of Boyacá, and selected 103 key plant species that, based on their uses, support the provision of ecosystem services in the páramos. We collated occurrence information for each key species and using a Mahalanobis distance approach we applied climate niche modelling for current and future conditions. Results We show an overall tendency of reduction in area for all ecosystem services under future climate conditions (mostly a loss of 10% but reaching up to a loss of 40%), but we observe also increases, and responses differ in intensity loss. Services such as Food for animals , Material and Medicinal , show a high range of changes that includes both positive and negative outcomes, while for Food for humans the responses are mostly substantially negative. Responses are less extreme than those projected for individual species but are often complex because a given ecosystem service is provided by several species. As the level of functional or ecological redundancy between species is not yet known, there is an urgency to expand our knowledge on páramos ecosystem services for more species. Our results are crucial for decision-makers, social and conservation organizations to support sustainable strategies to monitor and mitigate the potential consequences of climate change for human livelihoods in mountainous settings.

Adaptive Evolution Is Common in Rapid Evolutionary Radiations

Article

Full-text available

Sep 2019
CURR BIOL

One of the most long-standing and important mysteries in evolutionary biology is why biological diversity is so unevenly distributed across space and taxonomic lineages. Nowhere is this disparity more evident than in the multitude of rapid evolutionary radiations found on oceanic islands and mountain ranges across the globe [1-5]. The evolutionary processes driving these rapid diversification events remain unclear [6-8]. Recent genome-wide studies suggest that natural selection may be frequent during rapid evolutionary radiations, as inferred from work in cichlid fish [9], white-eye birds [10], new world lupins [11], and wild tomatoes [12]. However, whether frequent adaptive evolution is a general feature of rapid evolutionary radiations remains untested. Here we show that adaptive evolution is significantly more frequent in rapid evolutionary radiations compared to background levels in more slowly diversifying lineages. This result is consistent across a wide range of angiosperm lineages analyzed: 12 evolutionary radiations, which together comprise 1,377 described species, originating from some of the most biologically diverse systems on Earth. In addition, we find a significant negative correlation between population size and frequency of adaptive evolution in rapid evolutionary radiations. A possible explanation for this pattern is that more frequent adaptive evolution is at least partly driven by positive selection for advantageous mutations that compensate for the fixation of slightly deleterious mutations in smaller populations.

1054. Puya castellanosii L.B. Sm. a narrow Argentinian endemic species: Bromeliaceae

Article

Mar 2023

Puya castellanosii L.B. Sm., is illustrated and its relationships are discussed. Its habitat is described and illustrated, and suggestions for its cultivation are given. Seed was collected and it was raised and distributed by the plantsman Nori Pope who died in 2019.

Supplementary Material 13

Data

Full-text available

Feb 2019

Subfamilial Phylogenetic Relationships of the Bromeliaceae: Evidence from Chloroplast DNA Restriction Site Variation

Article

Full-text available

Jul 1990
SYST BOT

Restriction-site analyses of the chloroplast genomes of members of the Bromeliaceae were conducted to explore subfamilial phylogenetic relationships. Eighteen restriction-site mutations and one large (1.8-kb) length mutation were found to be variable at the family and subfamily levels. Cladistic analyses of these data supported the following hypotheses: 1) Bromelioideae and Pitcairnioideae are sister taxa and the latter may be paraphyletic; 2) Tillandsioideae s. str. and Bromelioideae are each monophyletic; 3) Tillandsioideae s. str. is the sister taxon of the remainder of the family; and 4) Glomeropitcairnia is phylogenetically distinct from the remainder of Tillandsioideae. These analyses suggest that the primarily terrestrial Pitcairnioideae are not basal in the family even though the terrestrial habit apparently is ancestral in Bromeliaceae. Chloroplast DNA data also support the contention that some morphological characters have evolved independently several times in this large, ecologically complex family.

Morphological Variation of Some Floral Features of the Subfamily Pitcairnioideae (Bromeliaceae) and Their Significance in Pollination Biology

Article

Full-text available

Mar 1988
Bot Gaz

Scanning electron and light microscopy observations of wet-preserved flowers of Bromeliaceae subfamily Pitcaimioideae yield new information on the stigma, petal scales, and septal nectaries. Variations of the stigma types are evident among several genera. The gross structural features of the stigma do not indicate definite pollination trends, but the shape of the lobes and papillae indicate a few specific modes. In pitcaimioid genera, petal scales, when functional, may aid in pollination by accumulating the nectar secreted from the ovary, thus facilitating its availability to the pollinator. Nectaries associated with the gynoecia usually display tripartite channels in the ovary septa. Some developmental changes of the channel structure and position of the ovary indicate three probable modes of nectar release from the gynoecia of the pitcairnioids: (I) through lateral grooves or openings, (2) partly through the apical orifices and partly through the dissolved areas of the spetal channels, and (3) exclusively through the apical orifices. Analysis of a wide range of floral features indicates that ornithophily, chiropterophily, and entomophily exist in different Pitcairnioideae lineages.

Methods for Computing Wagner Trees

Article

Full-text available

Mar 1970
Syst Zool

James Farris

Farris, J. S. (Biol. Set., State Univ., Stony Brook, N.Y.) 1970. Methods for computing Wagner Trees. Syst. Zool., 19:83-92.—The article derives some properties of Wagner Trees and Networks and describes computational procedures for Prim Networks, the Wagner Method, Rootless Wagner Method and optimization of hypothetical intermediates (HTUs).

The nature of cladistic data

Article

Jan 1987

Puya, the Pineapple's Andean ancestor

Article

Jan 1950

M.B. Foster

Repugnant and mature character concepts in phylogenetic inference: A temporal similarity and historical identity.

Conference Paper

Apr 2003

Arnold G. Kluge

The significance of "being similar" in the inference of species relationships is refuted once again (see also Hennig, 1966, Phylogenetic Systematics, Univ. of Illinois Press, Urbana, IL). Without merit is Rieppel and Kearney's (Biol. J. Linn. Soc., 2002, 75, 59-82) claim that submitting the relational property of topological similarity, their preferred definition of character, to falsifying tests of similarity benefits that kind of inference. Such a priori uses of similarity, in character analysis, are consistent with observational theory, where a character is defined intensionally in terms of immutable properties. However, the induced hypotheses that follow from this theory, not the deductive test that Rieppel and Kearney wanted, remain controversial, because their predictability is a consequence of circular reasoning, and their projectabality fails empirically from incongruent observation reports. Further, a category mistake is made when the abstract, similarity-defined, group of organisms is reified, as a part of history. In addition, Rieppel and Kearney failed to provide a special theory for similarity, which renders similarity scientifically repugnant (Quine, 1969, Ontological Relativity and Other Essays, Columbia Univ. Press, New York). A return to Hennig's (1966) evolutionary concept of evidence, as transformation series, is urged, and from which a testable character hypothesis can be formulated. There is no one operation for determining character states in this system-it can be anything that leads to the testable hypothesis of synapomorphy, as an historical identity relation. Character compatibility and conjunction, but not similarity, provide a priori tests in phylogenetic character analysis. In turn, the phylogenetic system of inference leads to explanations of homology, as historical identities, which exemplifies the goal of achieving a mature state of historical knowledge (not of Quine, 1969). Such maturity obtains from attempts to falsify hypotheses of species relationships with severely tested evidence, not from induction of "the" observation statement that Rieppel and Kearney sought to justify their true belief in a hypothesis of relationships.

Character States, Morphological Variation, and Phylogenetic Analysis: A Review

Article

Jul 1991

P. F. Stevens

The step in cladistic analysis that has received least attention is delimitation of character states, there usually being little justification for their delimitation. It is generally assumed that states of cladistic characters are discrete, even when variation is quantitative. I show here that a majority of the character states of obviously quantitative characters used in lower-level cladistic studies in botany over the last generation are ambiguous even when ingroup variation alone is analyzed. Consideration of variation in the outgroup may compromize either the states recognized in the ingroup and/or the polarity that they are subsequently assigned. Furthermore, many so-called qualitative characters are based on a quantitative phenomenological base filtered through the reified semantic discontinuities of botanical terminology; such characters face the problems of their more obviously quantitative relatives. Methods for delimiting states within quantitative characters are examined. Some produce gaps in the variation by redefining the character, scoring the intermediates in a distinctive fashion, performing phylogenetic analyses within the terminal taxa, or changing the hierarchical level at which the variation is evaluated. Others produce states by manipulation of the statistical properties of the variation of the ensemble of taxa being studied. These latter methods often allow greater resolution of the phylogeny, but at the cost of lowering the significance of the most parsimonious tree. The underlying assumptions of the two sets of methods are briefly analyzed. Problems manifest in the division of continuous variation into character states suggest a reappraisal of the early steps of cladistic analysis; in practice, character states often seem to be delimited in conjunction with developing ideas of the phylogeny, rather than in a step prior to a phylogenetic analysis. It is recommended that character states be delimited by carefully analyzed discontinuities (not necessarily absolute gaps) in the variation, attention having been paid to variation in the outgroup, and that "morphological" characters in general are assumed to be quantitative unless demonstrated otherwise. Explicit justification for the delimitation of character states should be given as a matter of course in all phylogenetic studies.

Notes on Bromeliaceae, XI

Article

Jan 1966

Lyman B. Smith

TNT (Tree analysis using New Technology)

Article

Jan 2000

Methods for Coding Variable Morphological Features for Numerical Taxonomic Analysis

Article

Sep 1985
Syst Zool

James W. Archie

Five methods of coding character differences for meristic or continuous variables when variability within taxa exists are discussed: (1) scaling by among-group variability; (2) scaling by within-group variability; (3) simple gap-coding; (4) homogeneous-subset coding; and (5) generalized gap-coding. Scaling by among-group variability is ineffective as it does not use within-group variability for coding purposes. Scaling by within-group variability does not effectively alleviate computational and theoretical problems associated with within-group character variability. The implications of gap-coding for character differentiation are examined, and the relationship between increasing sample size and decreasing information content of gap-coded characters is reviewed. The technique of generalized gap-coding is proposed to eliminate the problems apparent with simple gap-coding. Homogeneous-subset coding is developed in more detail than previously and compared to generalized gap-coding. The two techniques differ only in the critical values used to establish homogeneous or discriminant groups. These differences are most pronounced with unequal sample sizes for taxa. For moderate to large sample sizes, homogeneous-subset coding will detect more differentiation than generalized gap-coding for most choices of critical gap size. Seven data sets, representing a range of number of taxa and characters, as well as types of taxonomic units, were coded following the three procedures (3, 4, and 5, above), and phylogenetic trees were constructed. The results are discussed with regard to the amount of useful information retained by the coding procedures, the number of minimum-length trees found, and character consistency of the characters on the constructed trees. Consistency levels for all the coding methods were low and inversely related to the number of taxa present in the study. Finally, character-coding methods for phylogenetic analysis are discussed with regard to implications from the Kluge-Kerfoot phenomenon.