Botanical Journal of the Linnean Society, 2010, 162, 1–27. With 10 figures
A revision of Aechmea subgenus Macrochordion
(Bromeliaceae) based on phenetic analyses of
morphological variation
boj_1019
1..27
ANA PAULA GELLI DE FARIA1*, TÂNIA WENDT2 and GREGORY K. BROWN3
1
Departamento de Botânica, Universidade Federal de Juiz de Fora, ICB, Campus Universitário,
Bairro Martelos, 36036-900, Juiz de Fora-MG, Brazil
2
Departamento de Botânica, Universidade Federal do Rio de Janeiro, IB, CCS, Ilha do Fundão,
21941-590, Rio de Janeiro-RJ, Brazil
3
Department of Botany, University of Wyoming, 82071, Laramie, Wyoming, USA
Received 16 August 2009; accept for publication 9 November 2009
Considerable taxonomic confusion exists among species of Aechmea subgenus Macrochordion (Bromeliaceae), which
comprises the A. bromeliifolia complex. Cluster and principal components analyses were performed in order to
identify how many taxa exist in this complex and how they can be distinguished morphologically. Data for 16
morphological characters were scored from 38 selected specimens from different geographical regions, including
type specimens. Results from phenetic analyses, associated with observations in the field and of herbaria
exemplars, support the recognition of five species, one with two varieties (A. alba, A. bromeliifolia var. bromeliifolia, A. bromeliifolia var. albobracteata, A. lamarchei, A. maasii and A. triangularis). Five previously recognized
taxa are placed into synonymy: A. maculata and A. chlorophylla (both under A. lamarchei); A. pabstii (under A.
alba); A. kautskyana (under A. triangularis); and A. bromeliifolia var. angustispica (under A. bromeliifolia var.
bromeliifolia). This revision also includes a key to the species, descriptions, specimens examined, illustrations,
photographs, distribution maps and information on conservation status. © 2010 The Linnean Society of London,
Botanical Journal of the Linnean Society, 2010, 162, 1–27.
ADDITIONAL KEYWORDS: Bromelioideae – conservation status – morphology – multivariate analyses –
species complex – taxonomy.
INTRODUCTION
With more than 250 species (Luther, 2008), Aechmea
Ruiz & Pav. is the largest genus in Bromelioideae
(Bromeliaceae) and in the last monograph (Smith &
Downs, 1979) eight subgenera were recognized:
Aechmea, Chevaliera (Gaudich. ex. Beer) Baker, Lamprococcus Beer (Baker), Macrochordion de Vriese
(Baker), Ortgiesia (Regel) Mez, Platyaechmea (Baker)
Baker, Podaechmea Mez and Pothuava (Baker) Baker.
Nearly 70% of Aechmea spp. are from Brazil (Smith &
Downs, 1979; Luther & Sieff, 1994, 1997; Luther,
2001) and the Atlantic Forest is considered the centre
*Corresponding author. E-mail: [email protected]
of diversity for this genus and for most other genera
of Bromelioideae (Smith, 1934).
Infrageneric systematics of Aechmea is considered
artificial and poorly understood. The taxonomic
boundaries that separate Aechmea from other genera
of Bromelioideae are also vague and in need of revision. These problems were noted by early taxonomists
(Baker, 1879, 1889; Mez, 1891–94, 1896, 1934–1935)
and remain true in the most recent revision (Smith &
Downs, 1979). Discordant classifications in Aechmea
have emerged because monographers have stressed
only a few characters and knowledge of many potentially useful diagnostic data (e.g. floral, seed and fruit
morphology) is limited, these often being inaccessible
in herbarium material and, thus, variability is
poorly understood. Faria, Wendt & Brown (2004) also
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
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A. P. G. DE FARIA ET AL.
demonstraded that most characters traditionally
emphasized in taxonomic treatments of Aechmea
show considerable homoplasy and often fail to delimit
natural groups. After Smith & Downs (1979), taxonomic work within Aechmea has concerned mostly
description of new taxa, sometimes on the basis of
minor morphological variation. Published taxonomic
revisions dealing with Aechmea and other genera of
Bromelioideae are scarce (Wendt, 1997; Canela, Paz
& Wendt, 2003; de Sousa & Wendt, 2008).
Aechmea subgenus Macrochordion, the focus of this
study, is distinguished from other subgenera by the
simple, strobilate inflorescence with sessile, polystichous flowers, entire, unarmed, carinate floral bracts
densely covered by trichomes and by unarmed sepals.
However, the great morphological similarity shared
among the species makes delimitation difficult, particularly when diagnostic characters, especially reproductive, are poorly preserved or not mentioned on
herbarium specimen labels (e.g. calyx and corolla
colour). Furthermore, the available key for species
identification (Smith & Downs, 1979) includes some
characters of dubious interpretation, a consequence of
incomplete morphological descriptions.
At the beginning of this study, Macrochordion
included 11 taxa (A. alba Mez, A. bromeliifolia
var. bromeliifolia Rudge (Baker), A. bromeliifolia
var. albobracteata Philcox, A. bromeliifolia var.
angustispica Philcox, A. chlorophylla L.B.Sm., A.
kautskyana E.Pereira & L.B.Sm., A. lamarchei Mez,
A. maasii Gouda & W.Till, A. maculata L.B.Sm, A.
pabstii E.Pereira & Moutinho and A. triangularis
L.B.Sm.). The subgenus has a distribution entirely
restricted to the Brazilian Atlantic Forest, except for
A. bromeliifolia, which occurs from Central America
to Argentina.
Originally established at the generic level (de
Vriese, 1853), Macrochordion was later transferred
under Aechmea as a subgenus (Baker, 1889). This
infrageneric delimitation was kept in subsequent
revisions for Bromeliaceae (Mez, 1891–94, 1896,
1934–1935; Smith & Downs, 1979). Between the
establishment of Macrochordion and the publication
of the Smith and Downs monograph, many species
were described, synonymized or transferred to other
subgenera of Aechmea. A brief summary of the taxonomic history of Macrochordion includes: the description of A. alba, A. lamarchei and A. turbinocalyx in
Flora Braziliensis by Mez (1891–94); the four new
species described in a treatment of Bromeliaceae of
Brazil (Smith, 1955): A. chlorophylla, A. maculata, A.
nervata and A. triangularis; the description of A.
bromeliifolia var. albobracteata by Philcox (1974); and
the placement of 14 taxonomic synonyms in A. bromeliifolia var. bromeliifolia by Smith & Downs (1979).
At this point, Aechmea subgenus Macrochordion
included eight species. Later, four new taxa were
described for Macrochordion: A. pabstii (Pereira &
Moutinho, 1980), A. kautskyana (Pereira, 1980), A.
bromeliifolia var. angustispica (Philcox, 1992) and A.
maasii (Gouda & Till, 1997). Leme (1992) synonymized A. nervata under A. vanhoutteana (Van Houtte)
Mez, this last species belonging to Aechmea subgenus
Pothuava. Faria & Wendt (2004) proposed the reclassification of A. turbinocalyx from Aechmea subgenus
Macrochordion to subgenus Aechmea.
The goals of this paper were to conduct a detailed
taxonomic study of Aechmea subgenus Macrochordion. In addition to field and herbarium studies, we
also employed a phenetic analysis (cluster and principal components) as a tool to understand better the
patterns of morphological variation within the A. bromeliifolia complex and to show similarity relationships among taxa studied. Underutilized characters
received focus (e.g. floral structures) to improve
knowledge of the morphology of Bromelioideae.
MATERIAL AND METHODS
This taxonomic revision is based on herbarium collections, field observations, bibliographic information
and phenetic analyses of morphological data. Nearly
600 herbarium sheets were analysed from the following herbaria: ALCB, BHCB, CVRD, CEPEC, CESJ,
COR, CTES, GH, HB, HBR, HRB, HUEFS, HUFU,
IBGE, MBM, MBML, MO, NY, R, RB, RFA, SEL, SP,
SPF, TRIN, UB, VIC, VIES. Digital photographs from
G, K, LG, M, P, US and WU were also examined. Field
work was conducted in north-east and south-east
Brazil and collected specimens were deposited in
RFA. Some individuals studied in the field were cultivated at the greenhouse, Federal University of Rio
de Janeiro, in order to observe in more detail the
variability of selected characters. Live blooming specimens were photographed and fresh flowers were
preserved in 70% alcohol prior to examination. Morphological characters were documented for both
herbarium and liquid-preserved samples. When
possible, flowers from herbarium material were
rehydrated before analysis. The specific terminology
adopted for morphological descriptions follows
Radford (1986) and Smith & Downs (1979). Distribution and habitat data were taken from the herbarium
specimens and from the literature. Only nomenclatural changes published after Smith & Downs (1979)
are cited here. Conservation assessments were
generated using the IUCN red list category criteria
(IUCN, 2001).
The phenetic study included multivariate analyses
of cluster (CA) and principal component (PCA).
Cluster analysis was carried out to evaluate the relative similarities among taxa and principal component
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
AECHMEA SUBGENUS MACROCHORDION
analysis was conducted to identify those morphological traits that were most important in the differentiation of taxa. From all examined specimens, 38
provided sufficient and available data for the analyses, with intact and well-preserved vegetative and
reproductive structures. Each specimen was considered an operational taxonomic unit (OTU) and they
were selected to represent, as far as possible, the
entire geographical range and morphological variability present within each taxon. Data from A. bromeliifolia var. angustispica, A. maasii, A. maculata, A.
pabstii and A. kautskyana were obtained from the
type specimen only, through analysis of detailed
photos and/or direct examination of holotypes or isotypes, complemented with the original descriptions.
More than one OTU, including type specimens, was
analysed for A. alba, A. chlorophylla, A. lamarchei, A.
triangularis, A. bromeliifolia var. albobracteata and
A. bromeliifolia var. bromeliifolia. For this last taxon,
a representative specimen from the type locality was
sampled, because of the unavailability of the holotype, including suitable photographs.
Sixteen morphological characters (three vegetative
and 13 reproductive) were examined on each specimen and scored as binary or multistate (Table 1). The
continuous variable of leaf spine length was assigned
discrete states based on observed length gaps among
species. Vegetative characters were analysed from dry
specimens and the reproductive ones were documented from rehydrated or alcohol-preserved flowers.
For cluster analysis, pairwise relationships were estimated by the Manhattan Distance coefficient and the
resulting distance matrix displayed as a phenogram
using the UPGMA clustering method. The PCA was
based on a correlation matrix where only those
axes corresponding to components with eigenvalues
greater than 1.0 were extracted. Both data analyses
were carried out using the program Statistica 5.1
(StatSoft, 1998).
RESULTS AND DISCUSSION
Two major groups were found in cluster analysis, here
called A and B (Fig. 1). Group A contains A. bromeliifolia var. bromeliifolia, var. albobracteata, var. angustispica, A. triangularis and A. kautskyana, and was
further divided in two subgroups (C and D), thus
segregating A. triangularis and A. kaustyana from
the varieties of A. bromeliifolia (Fig. 1). Group B
comprises A. maculata, A. lamarchei, A. chlorophylla,
A. maasii, A. pabstii and A. alba and was divided in
subgroup E (A. chlorophylla, A. lamarchei and A.
maculata) and subgroup F, with this last segregating
A. maasii plus the remaining sampled OTUs identified as A. lamarchei from A. alba and A. pabstii
(subgroups G and H, respectively, Fig. 1).
In the PCA, the first three axes accounted for
75.43% of the total variance (45.78%, 19.22% and
10.43%, respectively; Table 2). The first component
was weighted heavily for leaf blade apex (2; numbers
in parentheses refer to characters in Table 1), floral
bract apex (7), sepal symmetry (9), sepal apex (10),
sepal fusion (11), petal shape (13), petal apex (14) and
fimbriate ligulae position (16). The second component
was weighted heavily for leaf spine length (2),
peduncle bract margins (5) sepal colour (12), petal
colour (15) and the third component for floral bract
texture (8). The same two major groups identified in
CA (groups A and B, Fig. 1) were also segregated in
the PCA (Fig. 2A), where A. bromeliifolia and variet-
Table 1. Qualitative characters and states used for the phenetic analysis
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
3
Habit: (0) epiphytic or rupicolous; (1) terrestrial
Leaf blade apex: (0) apiculate; (1) caudate
Leaf spine length: (0) up to 3 mm; (1) longer than 3 mm
Peduncle bract colour: (0) pink; (1) red; (2) white
Peduncle bract margins: (0) entire; (1) denticulate toward the apex
Peduncle bract arrangement and orientation: (0) subdense and divergent toward the apex of the peduncle
(1) imbricate and erect toward the apex of the peduncle
Floral bract apex: (0) truncate; (1) truncate–apiculate; (2) emarginated–apiculate; (3) obtuse–apiculate
Floral bract texture: (0) coriaceous with thin apex; (1) coriaceous throughout
Sepal symmetry: (0) symmetric to slightly asymmetric; (1) distinctly asymmetric
Sepal apex: (0) emarginate; (1) obtuse
Sepal fusion: (0) connate to the middle; (1) connate near the base
Sepal colour: (0) white to greenish white; (1) yellow to greenish yellow; (2) vinaceous
Petal shape: (0) spatulate; (1) lingulate
Petal apex: (0) obtuse; (1) emarginated
Petal colour: (0) white; (1) yellow to greenish yellow; (2) dark blue
Fimbriate ligulae position: (0) at distal end of the lateral folds; (1) approximately halfway along the lateral folds
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
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A. P. G. DE FARIA ET AL.
Figure 1. Phenogram obtained from cluster analysis of the Aechmea bromeliifolia complex using UPGMA. Names of each
OTU correspond to those recorded on herbarium sheets and are abbreviated as alb (A. alba); pab (A. pabstii); lam (A.
lamarchei); chl (A. chlorophylla); maa (A. maasii); mac (A. maculata); bbr (A. bromeliifolia var. bromeliifolia); ban (A.
bromeliifolia var. angustispica); bal (A. bromeliifolia var. albobracteata); kau (A. kautskyana) and tri (A. triangularis). *
indicates the type specimen or a specimen from the type locality. Main clusters discussed in the text are identified by the
letters A to H. Names on the left of the brackets correspond to those adopted after the taxonomic revision.
ies plus A. kautskyana and A. triangularis (group A)
can be characterized by leaf spines of 3 mm or longer,
symmetrical to slightly asymmetrical sepals, emarginated and connate to the middle, and spatulate and
emarginate petals. Likewise, the taxa of group B
(A. alba, A. chorophylla, A. lamarchei, A. maasii, A.
maculata and A. pabstii) can be characterized by leaf
spines of 1–3 mm, distinctly asymmetrical and obtuse
sepals, and lingulate and obtuse petals, with fimbriate ligulae about halfway along the lateral folds.
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
AECHMEA SUBGENUS MACROCHORDION
5
Table 2. Factor loadings and percentage of variance for the three principal components obtained from the 16 characters
analysed
Principal components
Characters
1
2
3
Habit
Leaf spine length
Leaf blade apex
Peduncle bract colour
Peduncle bract margins
Peduncle bract arrangement and orientation
Floral bract apex
Floral bract texture
Sepal symmetry
Sepal apex
Sepal fusion
Sepal colour
Petal shape
Petal apex
Petal colour
Fimbriate ligulae position
Variation explained (%)
-0.467813
0.264660
0.879458
-0.379908
-0.163165
0.242058
-0.741839
0.462271
-0.979068
-0.942595
-0.772111
0.413198
-0.948542
0.926107
0.503039
-0.802943
45.78
0.133173
0.700785
-0.092568
-0.403411
0.791543
-0.111028
0.428431
-0.408898
-0.057405
0.108021
-0.105821
0.780321
0.087313
0.084962
0.793290
0.355665
19.22
-0.450599
-0.465866
-0.015972
0.358292
-0.077763
0.492872
0.350477
-0.656441
0.101977
0.032188
-0.349294
0.242256
0.036852
-0.042609
0.147347
-0.317341
10.43
The analyses suggested little morphological differentiation between A. alba and A. pabstii (Figs 1, 2).
Petal colour was the variable that contributed most to
the separation of both species from other OTUs along
PC2 (Table 2). Apart from the white corolla, A. alba
and A. pabstii have morphologically identical floral
bracts, sepals and petals. Both taxa also share a
restricted and overlapping distribution in the south of
Bahia state. Close morphological similarity between
A. kautskyana and A. triangularis was also found
(Figs 1, 2B). Both taxa are endemic to dense ombrophile forest areas of Espírito Santo state. The
caudate–recurvate leaf apex and the dark-blue corolla
clearly distinguish these two species from others in
the subgenus. As suggested for A. alba and A. pabstii,
the multivariate analyses support field observations
and herbarium study, that A. kautskyana and A.
triangularis do not possess any significant morphological traits to justify their continued recognition as
distinct species.
With regard to the remaining taxa of the complex
(the yellow-flowered species), the analyses did not
support the previously recognized taxonomic status
of A. chlorophylla and A. maculata as distinct species
from A. lamarchei. Cluster analysis suggested a close
linkage among these taxa (group E, Fig. 1), distributed in rocky grasslands (campos rupestres), semideciduous and dense ombrophile forest habitats of
Minas Gerais and Espírito Santo states. Comparision
of PCs 1 and 3 revealed this same cluster of OTUs
(Fig. 2B). Floral bract texture was the variable that
contributed most to the separation of these taxa from
the other OTUs along PC3 (Table 2). In fact, A. chlorophylla and A. maculata resemble A. lamarchei in
several reproductive characters, including those here
considered diagnostic for this last species (floral
bracts ovate to widely ovate, coriaceous with thin
apex). High similarity was also indicated between A.
maasii and specimens from Espírito Santo and Rio de
Janeiro states, distributed in sandy coastal plains
(restingas) and tableland forests (florestas de tabuleiros) (group G, Figs 1, 2B). Until this study, A.
maasii was practically only known from the type
locality, as most exemplars in herbarium collections
had been identified under miscellaneous names (e.g.
A. lamarchei, A. bromeliifolia and A. chlorophylla).
Aechmea maasii resembles A. alba and A. lamarchei
in many morphological characters. This is illustrated
in the phenogram produced by CA (Fig. 1) and by the
intermediate position of this species on the scatterplot of PCs 1 vs. 3 (Fig. 2B). In addition to the
distribution and habitat, A. maasii can be characterized by the coriaceous, truncate–apiculate and
densely floccose floral bracts. The analyses also suggested that A. bromeliifolia var. angustispica has a
greater morphological affinity with the typical variety
than with A. bromeliifolia var. albobracteata (Fig. 1).
Variety angustispica is based on differences of leaf
indument and inflorescence shape (Philcox, 1992). In
Macrochordion, we observed that these characters
show considerable plasticity under field conditions,
within the same individual (leaf indument) or among
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
6
A. P. G. DE FARIA ET AL.
Figure 2. Principal component analysis (PCA) scatterplots of the first two axes (A) and the first and third axes (B) based
on 16 morphological characters of the Aechmea bromeliifolia complex taxa. Groups A, B, C, D, E, G and H are the same
as in Figure 1. Names for each symbol correspond to those adopted after the taxonomic revision.
individuals of a same population (inflorescence
shape). Aechmea bromeliifolia var. angustispica is
only known from the type locality, a place also with
numerous records for var. bromeliifolia. For A. bromeliifolia var. albobracteata, Philcox (1974) stressed
the white peduncle bracts as a distinctive character,
contrasting with the pink bracts of the typical
variety. We also observed other clear differences, such
as green peduncle and floral bracts compared with
these being vinaceous to dark purple in the typical
variety. Field observations and the herbarium study
indicated a sympatric occurrence of both taxa in
some areas of south-east and central Brazil.
However, A. bromeliifolia var. albobracteata assumes
its geographical integrity along the rest of its distribution in south of Brazil, Argentina and Paraguay.
Unlike A. bromeliifolia var. angustispica, the results
indicate that the maintenance of the status of A.
bromeliifolia var. albobracteata as a distinct variety
seems appropriate.
This study provided the first comprehensive analysis of the A. bromeliifolia complex. It supports the
recognition of six distinct taxa for Aechmea subgenus
Macrochordion: A. alba, A. bromeliifolia var. bromeliifolia, A. bromeliifolia var. albobracteata, A. lamarchei, A. maasii and A. triangularis, that were defined
on the basis of their morphological discontinuities,
following the phenetic species concept (Sneath, 1976).
Statistical analyses were important for evaluating the
status of some ignored or underutilized floral characters (e.g. sepal and petal morphology). Multivariate
analyses were also conducted to better understand
the delimitation of others species complexes in Bromeliaceae, as in Pitcairnia L’Hér. (Wendt et al., 2000)
and Vriesea Lindl. (Costa, Rodrigues & Wanderley,
2009). Future approaches dealing with reproductive
biology, population genetic data and phylogenetic
studies will be interesting to provide knowledge about
taxa of Aechmea subgenus Macrochordion in the light
of other species concepts.
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
AECHMEA SUBGENUS MACROCHORDION
KEY
TO THE TAXA OF
AECHMEA
SUBGENUS
7
MACROCHORDION
1. Leaf blades with spines 1–3 mm long; sepals distinctly asymmetric, apex obtuse; petals lingulate, apex obtuse ..
..................................................................................................................................................... 2
1′. Leaf blades with spines 3 mm or longer; sepals symmetric to slightly asymmetric, connate to the middle, apex
emarginate; petals spatulate, apex emarginate.......................................................................................4
2. Floral bracts ovate to widely ovate, coriaceous with thin apex, obtuse or emarginated, apiculate......................
.................................................................................................................................. 3. A. lamarchei
2′. Floral bracts depressed ovate, coriaceous, apex truncate, apiculate............................................................3
3. Calyx white or greenish white; corolla white. East of Brazil (Bahia and Minas Gerais) ..................... 1. A. alba
3′. Calyx yellow, yellow greenish or reddish; corolla yellow. East of Brazil (Espírito Santo and Rio de Janeiro).......
...................................................................................................................................... 4. A. maasii
4. Flowers 1.6–2 cm long; calyx vinaceous; corolla dark blue; fimbriate ligulae about halfway along the lateral folds.
East of Brazil (Espírito Santo).......................................................................................5. A. triangularis
4′. Flowers 1.4–1.6 cm long; calyx green or yellow greenish; corolla yellow or yellow greenish; fimbriate ligulae at
distal end of the lateral folds. Central America to Argentina ....................................................................5
5. Peduncle of the inflorescence and floral bracts vinaceous or dark purpureous; peduncle bracts pink. Central
America to south-east of Brazil .......................................................... 2.1. A. bromeliifolia var. bromeliifolia
5′. Peduncle of the inflorescence and floral bracts green, peduncle bracts white. Central, south-east and south Brazil,
Argentina, Paraguay ....................................................................... 2.2. A. bromeliifolia var. albobracteata
TAXONOMIC TREATMENT
AECHMEA SUBGENUS MACROCHORDION (DE VRIESE)
BAKER, Hand. Bromel. 34. 1889.
Type: Bromelia tinctoria Mart. [= Aechmea bromeliifolia (Rudge) Baker].
Basionym: Macrochordion de Vriese, Jaarb. Kon. Ned.
Maatsch. Tuinb. 1853: 14. 1853.
= ‘Macrochordium’ de Vriese in Beer, Fam. Bromel.
22: 145. 1857.
HERBS, epiphytic, rupicolous or terrestrial.
ROSETTES forming a tank, funnelform to tubulose,
with numerous leaves. LEAF SHEATHS orbicular,
narrowly elliptic to transversely elliptic, entire. LEAF
BLADES linear, lanceolate or linear–triangular,
sparsely to densely serrate, apex obtuse, acute or
caudate, usually apiculate. INFLORESCENCE simple,
forming a cylindrical compact spike; peduncle thin,
erect, green, reddish, vinaceous or dark purple, floccose or lanate; peduncle bracts alternate, imbricate or
subdense, erect or divergent toward the apex of the
peduncle, chartaceous, entire or denticulate toward
the apex, pink, red or white, cymbiform, apex acute
or acuminate. FLOWERS hermaphroditic, sessile,
polystichous, congested. FLORAL BRACT bicarinate
(the lowest) or tricarinate, having a 3-dimensional
form with the flowers in between, entire, coriaceous
throughout or thin toward the apex, white, green,
yellow greenish, reddish, purplish, castaneous or vinaceous, white floccose, appressed lepidote or lanate,
ovate, widely ovate or depressed ovate, shorter to
about equalling the sepal length, apex obtuse, emarginate or truncate, muticous or apiculate. SEPALS
symmetric to distinctly asymmetric, with a lateral
membranaceous wing, connate near the base or to the
middle, white, yellow, green, reddish or vinaceous,
white lanuginose or covered with white appressed
scales, apex obtuse or emarginate, muticous or
minutely apiculate. PETALS lingulate or spatulate,
yellow, yellow greenish, white or dark blue, turning
black after the anthesis, bearing a pair of appendages
on the adaxial surface formed by two lateral folds and
two fimbriate ligulae implanted on the lateral folds,
apex obtuse or emarginate, erect or slightly divergent.
STAMENS with antipetalous filaments adnate to the
petals and the antisepalous ones free, anthers dorsifixed. STIGMA conduplicate–spiralized. OVARY inferior,
epigynous tube short or conspicuous, placentation
axial, ovules caudate. FRUIT baccate, globose, greenish
or red; seeds many, light pink or pale brown, caudate.
1. Aechmea alba Mez, in Martius, Eichler & Urbain,
Fl. Bras. 3(3): 375. 1892. (Figs 3A–F, 10A, B)
Type: Brazil, Bahia, without specific locality, 1834,
Blanchet 2276 (Holotype G! photo, Isotype G! photo).
= Macrochordion alba (Mez) L.B.Sm. & W.J.Kress,
Phytologia 66(1): 77. 1989. H. E. Luther & E. Sieff,
Selbyana 15(1): 65. 1994, pro syn.
= Aechmea pabstii E.Pereira & Moutinho, Bradea 3:
86–7, 91, 95. 1980. Type: Brazil, Bahia, Porto Seguro,
estrada para Arraial de Nossa Senhora D’Ajuda,
5.ii.1980, Moutinho 59 (Holotype HB!). Paratype:
Bahia, Porto Seguro, Fazenda Brasília, 6.ii.1980,
Moutinho 68 (HB!), syn. nov.
= Macrochordion pabstii (E.Pereira & Moutinho)
L.B.Sm. & W.J.Kress, Phytologia 66(1): 77. 1989, syn.
nov.
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
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A. P. G. DE FARIA ET AL.
Figure 3. Aechmea alba. A, habit. B, detail of leaf margin. C, flower with floral bract. D, floral bract. E, sepals. F, petal
and stamens. A, B (Faria et al. 140). C–F (Faria et al. 139).
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
AECHMEA SUBGENUS MACROCHORDION
9
Figure 4. Distribution of (A) Aechmea alba, (B) A. lamarchei, (C) A. maasii and (D) A. triangularis.
Description: HERB epiphyte or terrestrial, 24.5–
47.5 cm high. ROSETTE funnelform or wide-funnelform, with 15–25 leaves. LEAF SHEATHS 10–15 ¥
4–9 cm, orbicular or elliptic, adaxial surface vinaceous, abaxial surface green. LEAF BLADES 20–55 ¥
2–5 cm, linear, concolorous green, apex obtuse or
acute, apiculate, margins serrate; spines castaneous,
1–3 mm long. INFLORESCENCE with spike 3–6 ¥
2–4 cm; peduncle green, 18.5–42 cm long, white floccose; peduncle bracts imbricate or subdense, divergent toward the apex of the peduncle, entire, red,
5.5–11 ¥ 1.5–3 cm, apex acuminate. FLOWERS 1.8–
2.1 cm long. FLORAL BRACTS 8–12 ¥ 11–17 mm,
depressed ovate, shorter than the sepals, coriaceous,
white, green or yellow greenish with red apex and
margins, white floccose or appressed lepidote, apex
truncate, apiculate. SEPALS 8–12 ¥ 5–7 mm, distinctly
asymmetric, connate near the base, white or white
greenish, covered with white appressed scales, apex
obtuse, muticous or minutely apiculate. PETALS
14–17 ¥ 4 mm, lingulate, white, apex obtuse, erect;
fimbriate ligulae about halfway along the lateral
folds. STAMENS with filaments 12–15 mm long;
anthers c. 5 mm long. STIGMA c. 2 mm long. OVARY
c. 4 mm long, bearing a short epigynous tube; ovules
0.5–1 mm long. FRUIT greenish; seeds c. 4 mm long,
pale brown.
Distribution and habitat: A. alba is almost restricted
to the south of Bahia state, being recorded at lower
frequency in boundary areas with Minas Gerais state
(Fig. 4A). It grows at 5–700 m altitude, in herbaceous,
shrubby and wooded restingas, dense ombrophile
forest and florestas de tabuleiros.
Conservation status: Near Threatened (IUCN, 2001).
Specimens examined: BRAZIL, BAHIA – BELMONTE:
Estação Experimental Gregório Bondar, 16.v.1979,
Silva et al. 365 (CEPEC); id., 16°08′S 39°15′W,
12.v.1993, Thomas et al. 9892 (CEPEC, HUEFS, NY).
CARAVELAS, rod. BR 418, 16 km do entroncamento
com a BA 001, 18.iii.1978, Mori et al. 9660 (CEPEC);
Guaratinga to São Paulinho, BA km 10, 29.iii.1973,
Pinheiro 2040 (CEPEC). ILHÉUS, c. 7 km na estrada
de Olivença–Vila Brazil, 30.v.1991, Carvalho et al.
3292 (CEPEC); ITAMARAJÚ: Rod. para F. São José de
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A. P. G. DE FARIA ET AL.
Baixo, 28.i.1972, Pinheiro 1792 (CEPEC); id., Rod.
Itamarajú–Teixeira de Freitas, 3 km from Itamarajú,
Fazenda Chapadão, 3.xi. 1983, Callejas et al. 1631
(CEPEC). NOVA VIÇOSA: 22.vii.1979, Martinelli 6025
(RB); id., 10.iv.1984, Hatschbach 47800 (MBM).
PORTO SEGURO: BR 05, km 5, 19.vi.1962, Duarte 6782
(HB, HBR, RB); id., in cultivation on Marie Selby
Botanical Gardens, 4.x.1996, Anderson 48 (SEL); id.,
1.ix.1999, Luther s.n. (SEL 081043); id., Parque
Nacional de Monte Pascoal, 25.vi.1967, Castellanos
25519, 26515 (CEPEC); id., 21.iii.1968, Vinha &
Santos 90 (CEPEC); id., 15°15′53″S 40°34′29″W,
25.iii.1996, Thomas et al. 11161 (CEPEC, NY); id.,
12°52′02″S 37°24′54″W, 5.ii.1999, Thomas et al. 11979
(CEPEC); id., Rod. para Caraíva, 6.iv.2003, Faria
et al. 143 (RFA); id., Rod. para Sta Cruz de Cabrália,
km 7, 20.iv.1982, Carvalho et al. 1227 (CEPEC, HRB,
MBM); id., Reserva Biológica Pau–Brazil, 4.viii.1973,
Hage 69 (CEPEC); id., 16°25′S 39°12′W, 19.iii.1974,
Harley 17183, 17186 (CEPEC, RB); id., 19.iv. 1982,
Carvalho et al. 1187 (CEPEC); id., 16°23′27″S
39°10′48″W, 4.iv.2003, Faria et al. 139 (RFA). POTIRAGUÁ: Road to Potiraguá, 26.3 km da rodovia BR-415,
15°22′38″S 39°58′28″W, 2.xi.2000, Jardim et al. 3139
(CEPEC, NY); id., Road Una–Olivença, 16.vi.1971,
Pinheiro 1379 (CEPEC); id., Rod. Porto Seguro–
Eunápolis, km 12, 4.iv.1972, Eupunino 256 (CEPEC);
id., km 8, 26.xi.1975, Hage 138 (CEPEC, RB). STa
CRUZ DE CABRÁLIA, 6.iv.1979, Mori et al. 11692
(CEPEC); id., estrada para Porto Seguro, 19.iii.1968,
Vinha & Santos 64 (CEPEC); id., 21.v.1985, Martinelli et al. 11135 (NY, RB); id., estrada para Santo
André, 16°15′S 39°1′30″W, 17.vi.1980, Silva & Brito
878 (CEPEC); id., 16°7′90″S 38°59′39″W, 5.iv.2003,
Faria et al. 140 (RFA), id., 16°14′43″S 39°1′50″W,
5.iv.2003, Faria et al. 141, 142 (RFA); id., próximo ao
lixão, 7.iv. 2003, Faria et al. 145 (RFA); id., Reserva
Biológica do Pau Brazil, 17.ix.1971, Santos 1953
(CEPEC); id., rodovia Estação Ecológica Pau Brazil–
Sta Cruz, 5–7 km NE da Estação, 16°23′S 39°08′W,
5.vii.1979, Mori et al. 12088 (ALCB, CEPEC). STa
TERESINHA: in cultivation on Marie Selby Botanical
Gardens, 10.iii.1997, Berg s.n. (SEL 076901); id.,
6.v.2002, Luther s.n. (SEL 086394). TEIXEIRA DE
FREITAS: Vale do Rio Alcobaça, 12.v.1971, Santos
1624 (CEPEC). UNA, estrada Una–Olivença, km 4,
12.viii.1994, Ramírez et al. 478 (CEPEC); id., Reserva
Biológica de Una, 2.iv.2003, Wendt et al. 478A (RFA);
id., in cultivation on UFRJ, 5.i.2004, Wendt et al. 462
(RFA); id., Reserva Biológica do Mico–Leão, entrada
km 16 da rodovia BR 001 Ihéus–Una, Fazenda
Dois de Julho, 15°09′S 39°05′W, 14.vii.1993, Jardim
et al. 156 (CEPEC); id., 20.vii.1996, Amorim 1847
(CEPEC); id., rodovia Olivença–Una, km 35, próximo
a Reserva Biológica do Mico Leão, 2.vi.1981, Hage &
Santos 819 (CEPEC, RB); id., 19.v.1985, Martinelli
et al. 11121 (NY, RB). MINAS GERAIS – SALTO DA
DIVISA: 3.vi.1990, Hatschbach 54114 (MBM);
id., Fazenda Santana, 16°04′16,2″S 40°03′19″W,
20.ii.2003, Lombardi et al. 5094 (BHCB); STa MARIA
DO SALTO: Fazenda Duas Barras, 16°24′40,7″S
40,2°50,5′W, 23.viii.2003, Lombardi et al. 5409 (RFA).
Notes: When Pereira & Moutinho (1980) described A.
pabstii, they proposed an affinity with A. lamarchei,
A. chlorophylla and A. triangularis. Even although
the white corolla was considered relevant for the
identification of this species, the authors disregarded
the fact that the same character was found in A. alba,
described almost 90 years earlier (Mez, 1891–94) for
the same region of A. pabstii. The fact that A. alba
was described as having petiolate leaves and that this
feature was stressed by Smith & Downs (1979) in the
taxonomic treatment of the subgenus, could explain
why, until now, A. alba and A. pabstii were considered
distinct species. We did not detect, however, the presence of petiolate leaves in the holotype or isotype of A.
alba or in other herbarium specimens examined in
this study. The phenetic analyses suggested a close
affinity between A. alba and A. pabstii. The comparision of type materials and original descriptions
indicate that they should be treated as synonyms.
Aechmea alba can be characterized by the coriaceous,
depressed ovate and truncate–apiculate floral bracts
(Fig. 3D) and, especially, by the white calyx and
corolla (Fig. 10A, B). It resembles A. lamarchei and A.
maasii in having leaf spines up to 3 mm long, obtuse
and distinctly asymmetric sepals and obtuse, lingulate petals with fimbriate ligulae about halfway along
the lateral folds (Fig. 3B, E, F). The species is well
represented in herbaria, where most of the exemplars
were previously erroneously identified as A. lamarchei or A. chlorophylla.
2. Aechmea bromeliifolia (Rudge) Baker, in Bentham
& Hooker f., Gen. pl. 3: 664. 1883.
Description: HERB epiphytic, rupicolous or terrestrial, 50.5–100 cm high. ROSETTE funnelform or
tubular, with 15–20 leaves. LEAF SHEATHS 13.5–
34.5 ¥ 4–15.5 cm, elliptic to narrow-elliptic. LEAF
BLADES 26.5–85.5 ¥ 3–8 cm, linear or lanceolate,
apex obtuse or acute, apiculate, margins serrate or
sparsely serrate. INFLORESCENCE with spike 4.5–
13 ¥ 2–4 cm; peduncle 40.5–90 cm long, white lanate;
peduncle bracts imbricate or subdense, erect or
divergent toward the apex of the peduncle, entire,
8–17.5 ¥ 1.5–2.5 cm, apex acuminate. FLOWERS 1.4–
1.6 cm long. FLORAL BRACTS 8–11 ¥ 14–16 mm,
depressed ovate, shorter than the sepals, coriaceous,
white lanate, apex truncate or truncate–emarginate,
muticous. SEPALS 5–8 ¥ 5–7 mm, symmetric to
slightly asymmetric, connate to the middle, white
lanuginose, apex emarginate, muticous. PETALS
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AECHMEA SUBGENUS MACROCHORDION
11–12 ¥ 4–5 mm, spatulate, apex emarginate, erect or
slightly divergent; fimbriate ligulae at distal end of
the lateral folds. STAMENS with filaments 6.5–8 mm
long; anthers 3–4 mm long. STIGMA c. 1 mm long.
OVARY c. 4 mm long, bearing a short epigynous tube;
ovules c. 0.5 mm long. FRUITS greenish; seeds c. 4 mm
long, pale brown.
2.1. Aechmea bromeliifolia var. bromeliifolia
(Figs 5A–E, 10C, D)
Type: Guiana Francesa, s.d., Martin s.n. (Holotype
BM! photo).
Basionym: Tillandsia bromeliaefolia Rudge, Pl.
Guian. 32, t. 50. 1807.
= Macrochordion bromeliifolia (Rudge) Beer in
L.B.Sm. & W.J.Kress, Phytologia 66(1): 77. 1989. H.
E. Luther & E. Sieff, Selbyana 15(1): 65. 1994, pro
syn.
= Aechmea lagenaria Mez, in Martius, Eichler &
Urbain, Fl. Bras. 3(3): 372. 1892. Type: Brazil,
viii.1884, Morren Hortus s.n. (Holotype LG! photo),
syn. nov.
= Aechmea bromeliifolia var. angustispica Philcox,
Kew Bull. 47(2): 268, 270. 1992. Type: Brazil, Bahia,
27.5 km south-east de Morro do Chapéu, on road to
Mundo Novo, i.1977, Storr 145 (Holotype CEPEC!,
Isotype K! photo), syn. nov.
Description: LEAF SHEATHS with adaxial surface vinaceous or purpureous, abaxial surface green or sometimes reddish. LEAF BLADES concolorous green or
concolorous reddish, sometimes with abaxial surface
reddish and adaxial surface green, margins serrate or
sparsely serrate; spines black or sometimes reddish,
0.3–1 cm long. INFLORESCENCE with peduncle vinaceous or dark purple; peduncle bracts pink. FLORAL
BRACTS vinaceous or purplish. SEPALS green or
yellow–greenish. PETALS yellow or yellow–greenish.
Distribution and habitat: Aechmea bromeliifolia var.
bromeliifolia occurs from Central America (Mexico,
Guatemala, El Salvador and Honduras), to northwest of South America (Guianas, Suriname, Trinidad
and Tobago, Venezuela, Colombia, Peru, Bolivia) and
Brazil (Maranhão, Ceará, Bahia, Minas Gerais,
São Paulo, Tocantins, Goiás, Mato Grosso, Amapá,
Roraima, Pará, Amazonas and Rondônia states and
in Distrito Federal (Fig. 6). It grows at 140–1700 m
altitude, in Amazonian campinas and terra firme
forests, semi-deciduous forests, savannas (cerrados),
rocky grasslands (campos rupestres) and caatinga
vegetation.
Conservation status: Least Concern (IUCN, 2001).
Specimens Examined: BOLIVIA, BENI – VACA DIEZ:
camino de Riberalta hacia Guayaramerín, 18°08′S
65°45′W, 15.viii.2000, Kromer & Acebey 1450 (SEL);
id., on road to Cachuela Esperanza, 11°05′S 65°50′W,
7.ix.1981, Solomon 6179 (MO, SEL). CHUQUISACA –
11
HERNANDO SILES: 10 km de Monteagudo a Padilla,
19°48′S 64°01′W, 1.vii.1995, Kessler et al. 4973 (SEL).
LA PAZ – FRANZ TAMAYO: Parque Nacional Madidi,
refugio Chalalán, campamento Estabon y alrededores, 14°27′S 67°56′W, 24.iv.2000, Kromer & Acebey
1100 (SEL). SUD YUNGAS: rio Bopi, San Bartolome,
near Calisaya, vii.1939, Krukoff 10248 (GH, NY); id.,
5 km de Chamaca a La Asunta, 16°13′S 67°13′W,
6.x.1995, Kessler et al. 5812 (SEL). PANDO – NICOLAS
SUÁREZ: junto a Puerto Rico, riberas del río Tahuamanu, 26.i.1983, Casas 8484 (NY). SANTA CRUZ –
CHIQUITOS: Santiago, 3.ix.1942, Cutler 7035 (GH).
ICHILO: 4 km south-west campamento Macuñucu,
17°44′S 63°35′, 27.ix.1996, Kessler et al. 8684 (SEL).
NUFLO DE CHÁVEZ: 17°47′S 63°11′W, 26.vii.1987, Nee
35769, 35771 (NY). VELASCO: campamento El
Refugio, 14°46′39″S 61°02′52″W, 15.x.1994, Gullén &
Choré 2354 (WU). TARIJA – Aniceto ARCE: Bermejo,
9 km west and road Sta Cruz–Samaipata, 3.viii.1982,
Till 145 (WU). BRAZIL, AMAZONAS – distrito
agropecuário da SUFRAMA, 80 km de Manaus,
02°26′S 59°48′W, 23.vi.1992, Nee 42834 (NY);
Manaus–Caracaraí road, km 130, Igarapé, Lages,
9.v.1974, Prance et al. 21054 (NY); Manaus–
Itacoatiara highway, km 202, near Rio Urubu,
19.xii.1966, Prance et al. 3736 (NY); rio Cuieras,
below mouth of rio Brancinho, 29.ix.1971, Prance
et al. 15038 (NY); id., 15.ix.1973, Prance et al. 17958
(NY); id., 02°41′S 60°19′W, 21.vi.1992, Mori & Gracie
22429 (NY); rio Cunhuá, Deni indian village, 06°43′S
66 47′W, 28.ix.1971, Prance et al. 16471 (NY);
Xiborem, rio Negro, 19.viii.1928, Luetzelburg 22013
(R). AMAPÁ – rio Araguari, road side forest between
Porto Platón and Macapá, 18.ix.1961, Pires et al.
51090 (NY). BAHIA – ABAÍRA: distrito de Catolés,
13°17′30″S 41°52′39″W, 19.ix.1999, Nunes et al. 75
(HUEFS). BARRA DE ESTIVA–ITUAÇU: 23.iii.1980,
Araújo 280 (HRB, RB). CAETÉ–AÇU: cachoeira da
Fumaça, 30.iii.1993, Esteves & Kameyana 2527 (SP).
CAETITÉ: on road to Brejinho das Ametistas, 14°07′S
42°30′W, 13.iv.1980, Harley et al. 21317 (CEPEC); id.,
14°04′49″S 42°30′50″W, 7.viii.1996, Carvalho et al.
6255 (CEPEC). IBIQUERA: 13°3′09″S 41°18′33″W,
22.vi.1978, Vaillant 64 (RB); Ibiquera–Cascavel,
trilha para o Rumo (Machobongo), entre Riachão e
Morro do Chapéu, 4.xi.1989, Ferreira 243 (HRB, RB).
ITIRUÇÚ–MARACÁS: km 29, Fazenda Contendas,
7.vii.1971, Pinheiro 1429 (CEPEC). ITUAÇÚ: arredores
do Morro da Mangabeira, 13°50′22″S 41°18′43″W,
20.vi.1987, Queiroz et al. 1628 (HUEFS). LENÇÓIS:
Serra da Chapadinha, 12°27′13″S 41°26′50″W,
30.vii.1994, Pereira et al. 328 (ALCB); id., 12°27′35″S
41°26′25″W, 27.x.1994, Carvalho et al. 1108 (ALCB).
MARACÁS: Fazenda Gameleira, rod. BA 250, trecho
Itiriçú–Maracás, km 25, 29.ii.1988, Silva et al. 2247
(CEPEC). MORRO DO CHAPÉU: estrada para Morrão,
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
12
A. P. G. DE FARIA ET AL.
Figure 5. Aechmea bromeliifolia var. bromeliifolia. A, habit. B, flower with floral bract. C, floral bract. D, sepals. E, petal
and stamens. (Faria 176).
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
AECHMEA SUBGENUS MACROCHORDION
13
Figure 6. Distribution of Aechmea bromeliifolia var. bromeliifolia and A. bromeliifolia var. albobracteata.
11°35′03″S 41°11′31″W, 5.viii.2001, Nonato et al. 975
(HUEFS); Morro do Ouro, 19.vii.1981, Giulietti et al.
1334 (SPF). MUCUGÊ: 3 km S de Mucugê, estrada
para Jussiapi, 13°00′S 41°24′W, 26.vii.1979, Mori
et al. 12649 (RB); Mucugê–Guiné, 5 km de Mucugê,
7.ix.1981, Furlan et al. 1958 (CTES, RB, SPF).
PALMEIRAS: Morro Pai Inácio, 14.ii.1989, Till 4042
(WU); id., 12°27′20″S 41°28′15″W, 27.xii.1994, Guedes
et al. 1386 (ALCB, CEPEC); id., 12°27′31″S
41°28′17″W, 24.iv.1995, Costa et al. 1763 (ALCB).
PIATÃ: Serra do Atalho, entre Cravada e Cravadinha,
13°07′S 41°54′W, 22.viii.1992, Ganev 940 (HUEFS).
RIO DE CONTAS: 9–11 km N de Rio de Contas, estrada
para o povoado Mato Grosso, 13°32′S 41°46′W,
20.vii.1979, Mori et al. 12331 (CEPEC). SÃO FELIPE:
16.vii.1978, Heringer et al. 17116 (IBGE). SEABRA:
Morro do Pai Inácio, N da BR 242, 45 km de Seabra,
14.ii. 1989, Till 4040 (WU); Serra do Sincorá, 20 km
west of Barra da Estiva, 13°35′S 41°27′W, 22.iii.1980,
Harley et al. 20721 (CEPEC, SPF); vicinity of Toca da
Onça, vi.1915, Rose & Russell 20108 (GH photo).
CEARÁ – BATURITÉ: Serra do Baturité, 28.i.1974,
Reitz 7556 (HBR). CRATO: 12 km south-west of
Crato, on road to Exú, Serra do Araripe, 7°14′55″S
39°29′53″W, 30.vii.1997, Thomas et al. 11693
(CEPEC); Jaquara, Serra do Araripe, 11.ii.1935, Luetzelburg 26461 (GH). UBAJARA: Parque Nacional de
Ubajara, 03°49′S 40°53′W, 12.viii.1998, Martinelli
et al. 15055 (RB). DISTRITO FEDERAL – BRASÍLIA:
APA de Cafuninga, 15°33′S 48°06′W, 11.ix.1990,
Azevedo & Alvarenga 930 (IBGE); c. 10 km east of
Brasília, near Sobradinho, 1.x.1965, Irwin et al. 8861
(NY, UB); c. 30 km north-east of Brasília, 14.v.1966,
Irwin et al. 15835 (NY, UB); in cultivation on Estação
Experimental de Biologia da UNB, 26.ix.1973,
without collector (HB, UB); east side of Lagoa
Paranoá, 17.ix.1965, Irwin et al. 8401 (NY); entre
Brasília e Niquelândia, 10.v.1963, Pires et al. 9717
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A. P. G. DE FARIA ET AL.
(UB); Fazenda Água Limpa (UNB field station, near
Vargem Bonita), 21.x.1976, Ratter et al. 3819 (UB);
pastagem perto do ribeirão Água Doce ou Cafuninga,
15°31′S 47°58′W, 17.vii.1980, Kirkbride & Kirkbride
1315 (UB); Reserva Ecológica do Roncador, picada
R-5, 13.ix.1977, Heringer et al. 63 (IBGE); id., picada
R-4, 17.x.1977, Heringer et al. 227 (IBGE); Reserva
Ecológica do IBGE, 15°54′54″S 47°53′46″W, 22.v.1989,
Alvarenga 268 (HRB, IBGE, RB, SP). GOIÁS – ALTO
PARAÍSO DE GOIÁS: Chapada dos Veadeiros,
17.xii.1967, Duarte 10629 (HB); c. 22 km N de Alto
Paraíso, 22.iii.1971, Irwin et al. 32963 (NY, UB);
c. 7 km by road south Terezina, 17.iii.1973, Anderson
et al. 7347 (NY, UB); c. 52 km W de Alto Paraíso,
estrada para Niquelândia, 15.x.1980, Martinelli et al.
7537 (RB); c. 56 km N de Alto Paraíso de Goiás,
30.xi.1988, Kral et al. 75765 (SP); c. 23 km de Alto
Paraíso em direção a Teresina de Goiás, PN Chapada
dos Veadeiros, 13°56′39″S 47°29′38″W, 12.xi.1996,
Silva & Santos 3210 (IBGE, SP); Chapada dos Veadeiros, 6.v.1966, Heringer 11096 (HB); id., 24.ix.1967,
Haas & Belém 275 (HB); id., 17.iii.1969, Irwin et al.
24633 (NY, UB); id., iii.1969, Irwin et al. 24634 (NY,
UB); estrada Alto Paraíso–Terezina, 7 km após Alto
Paraíso, 14°06′02″S 47°31′28″W, 7.xii.1988, Neto 129
(IBGE). ARAÇÚ: Mato Grosso de Goiás, Fazenda São
José, 13.vii.1972, Leinig 520 (HB). COCALZINHO DE
GOIÁS: estrada Cocalzinho–Pirenópolis, próximo a
Três Picos, Serra dos Pirineus, 28.v.1998, Forzza et al.
908 (SPF). CRISTALINA: RPPN Linda Serra dos
Topázios, 16°45′S 47°40′W, 23.vi.1997, Proença &
Oliveira 1784 (UB); Filadélfia, Serra da Mamoneira,
5.viii.1964, Prance & Silva 58571 (GH, NY, UB).
GUARANI DE GOIÁS: estrada Posse–Guarani de Goiás,
ramal à direita, 2 km antes da divisa com a Bahia,
13°58′S 46°10′W, 29.vii.2000, Forzza et al. 1568 (SPF).
MOSSÂMEDES: Serra Dourada, cerca de 6 km Nordeste
de Mossâmedes, 16°04′S 50°11′W, 7.ii.1980, Kirkbride
3319 (UB). NIQUELÂNDIA: entrada a direita do km 06,
Niquelândia–Companhia de Níquel de Tocantins,
14°25′02″S 48°26′11″W, 29.vi.1996, Azevedo et al.
1046 (IBGE). POSSE: Posse para Bahia, 16.vi.1979,
Pereira 22 (HB); road to Cavalcante, 14°S 47°W,
22.x.1965, Irwin et al. 9516 (NY, MO, UB); rodovia
GO-118, cerca 14 km N de Alto Paraíso de Goiás,
21.xi.1987, Mamede et al. 87 (SP); Serra Dourada,
1969, Rizzo 4092, 4094 (RB); id., 10.v.1973, Anderson
et al. 10014 (NY, UB). SERRA DOS PIRINEUS: c. 18 km
E de Pirenópolis, 18.i.1972, Irwin et al. 34489
(NY, UB). MARANHÃO – STA LUZIA: estrada Duas
Barracas–Caldeirão, 7.x.1987, Monteiro et al. 20
(HRB, RB). MATO GROSSO – vii.1894, Kuntze s.n.
(NY). ALTO TAQUARÍ: c. 25 km sudeste da cidade,
estrada para Fazendas Córrego de Laje, 17°50′S
53°17′W, 21.ii.1996, Silva et al. 3010 (MBM).
ARIPUANÃ: rio Alto Juruena, vii.1962, Mee s.n. (SP
69028); Camizão, 24.ix.1940, Foster 1090 (GH); Guia,
18.v.1894, Lindman A3521 (GH photo). SÃO FÉLIX:
Fazenda Patizal, 12°49′S 51°46′W, 26.vii.1968, Richards 6523 (UB). SÃO LUIZ DE CÁCERES: ix.1908,
Hoehne 383 (R); id., Hoehne 439 (GH photo, R); id.,
ix.1911, Hoehne 4723, 4724 (R). MINAS GERAIS –
ARCOS: in cultivation on UFRJ, 5.ix.2003, Faria 176
(RFA); id., 16.x.2003, Faria 179 (RFA). BELO HORIZONTE: c. 140 km north of Belo Horizonte, Serra do
Cipó, 19.ii.1968, Irwin et al. 20487 (NY, UB); id.,
campus da UFMG, 2.viii.1998, Marques & Formiga
s.n. (BHCB 42945); id., dependências internas da
UFMG, 23.ix.1998, Marques & Formigas s.n. (BHCB
43379); id., Estação Ecológica da UFMG, 17.viii.1993,
Neto & França 858 (BHCB, US); id., Horto ICB,
Department de Botânica da UFMG, 23.ix.1998,
Marques & Formiga s.n. (BHCB 43372). BRUMADINHO: Serra da Calçada, retiro das Pedras, 20°08′S
44°13′W, 5.v.1990, Martins 398 (SPF); id., 20°05′35″S
43°59′01″W, 11.viii.2001, Viana 118 (BHCB). CALDAS:
1869, Regnell III 1255 (US); id., Regnell III 1726 (GH
photo). CARMO DO RIO CLARO: Fazenda Novo Horizonte, 26.viii. 1961, Andrade & Emmerich 937 (HB,
R); id., 2.ix.1961, Andrade & Emmerich 998 (R).
CATAS ALTAS: Serra do Caraça, trilha para o Pico do
Inficionado, 10.i.2000, Vasconcelos s.n. (BHCB 52560).
CONCEIÇÃO DO MATO DENTRO: Parque Natural
Municipal do Ribeirão do Campo, 19°06′12″S
43°34′28″W, 7.vii.2002, Mota & Viana 1833 (BHCB).
CONGONHAS: 8.viii.1980, Gurken & Gurken 16 (HB).
CONSELHEIRO MATTA: vi.1934, Brade 13971 (RB).
CRISTÁLIA: Serra do Batieiro, 14.ix.1991, Carvalho
553 (BHCB). CURVELO: estrada Belo Horizonte–
Brasília, cerca 15 km de Curvelo, 26.vi.1972, Braga
2537 (RB). DIAMANTINA: iv.1892, Schwacke 8412 (RB);
id., 1.ii.1947, Egler s.n. (RB 59651); id., 3.vi.1955,
Pereira 1675 (HB, RB); id., 2.iv.1957, Pereira & Pabst
2789 (RB); id., 3.viii.1973, Seidel 657 (HB); id.,
20.i.1972, Hatschbasch et al. 29045 (MBM); id.,
estrada para Biri-Biri, afloramento acima da cachoeira Sentinela, 14.vii.1996, Parra et al. 108 (SPF);
id., estrada para Conselheiro Mata, km 189,
2.viii.1985, Pirani et al. 7928 (SPF); id., estrada
Diamantina–Mendanha, km 578, 19.iii.1993, Esteves
& Kameyama 2473 (SP); id., km 572, 18°07′S
43°35′W, 26.ix.1994, Splett 699 (UB); id., estrada
Diamantina–Turmalina, 33 km de Diamantina,
14.v.1979, Martinelli 5975 (RB); id., road from Sopa to
São João da Chapada, 1 km north-west of Sopa,
29.i.1995, Till et al. 11048 (WU); id., Serra do Espinhaço, 3.ii.1972, Anderson et al. 35260 (UB). ENTRE
RIOS DE MINAS: 5.ix.1970, Krieger 9114 (CESJ).
GOUVÊIA: 30 km by road south-west of Gouvêia, at
km 60 on road to Curvelo, Serra do Espinhaço,
11.iv.1973, Anderson et al. 8634 (MO, NY, UB, US).
GRÃO MOGOL: campo rupestre atrás da cidade,
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AECHMEA SUBGENUS MACROCHORDION
13.iv.1981, Cordeiro et al. 792 (RB, SP, SPF); id.,
Córrego do Pasto, 21.x.1978, Hatschbasch & Kasper
41618 (MBM); id., em direção ao nordeste da cidade,
16°32′S 42°55′W, 22.v.1982, Giulietti et al. 3493 (RB,
SP, SPF); id., estrada para Barão, 14.v.1998, Forzza
et al. 805 (SPF); id., Jambeiro, 7 km de Grão Mogol,
5.ix.1985, Cavalcanti et al. 8550 (SPF); id., próximo a
saída da estrada para Francisco Sá, 7.vii.1986,
Meguro et al. 8999 (SPF); id., road to Cristália, Serra
do Espinhaço, 20.ii.1969, Irwin et al. 23608 (NY, UB,
US); id., trilha da Tropa, 11.xii.1989, Pirani et al.
12482 (SPF); id., Vale do Rio das Mortes, oeste da
cidade, 24.vii.1986, Mello-Silva et al. 9945 (RB, SPF).
INDIANÓPOLIS: parcela 5 UHE Miranda, 20.viii.1998,
Vasconcelos s.n. (BHCB 42869). ITABIRITO: Pico do
Itabirito, Serra dos Inconfidentes, 2.vii.1993, Teixeira
s.n. (BHCB 24107). ITUIUTABA: 16.ix.1956, Macedo
2591 (US). JABOTICATUBAS: Lapinha de Cima,
próximo a RPPN Ermo de Minas, 28.viii.2003, Faria
& Versieux 168, 169, 170 (RFA); id., km 115 rodovia
Lagoa Santa–Conceição do Mato Dentro–Diamantina,
16.viii.1979, Wanderley 5688 (SP); id., Serra do Cipó,
21.v.1975, Reitz 7862 (HBR); id., 6.ix.1976, Menezes
819 (SP); id., 5.viii.1980, Wanderley et al. 218 (SP).
LAGOA SANTA: APA Carste de Lagoa Santa, 1995,
Brina s.n. (BHCB). MARIANA: Área da SAMARCO,
15.viii.2000, Brina s.n. (BHCB 60049). MONTE
BELO: Fazenda Queimada Grande, 21°24′S 46°17′W,
7.ix.1987, Gentry 59138 (MO). OURO BRANCO:
28.xii.1937, Castellanos 20585 (CTES, GH); id., cultivada por R. A. Kautsky, 26.vi.1986, Kautsky 427 (RB);
id., Serra do Ouro Branco, 20°28′S, 43°41′W,
12.v.1990, Arbo et al. 3989 (CTES, MBM, SPF); id.,
29.viii.1996, Filho s.n. (BHCB 32937, 32938); id.,
19.ix.1998, Marques et al. s.n. (BHCB 43368, 43369,
43371, 43375, 43376, 43377); id., 25.iv.2001, Silva 157
(VIC); id., 9.vi.2001, Silva 170 (VIC); id., 10.vi.2002,
Paula et al. s.n. (VIC 27384); id., 28.vi.2002, Paula
et al. s.n. (VIC 27383). OURO PRETO: 20.vii.1894,
Schwacke 10557 (RB); id., x.1896, Silveira 1846
(R); id., Lavras Novas, 23.ii.2002, Paula et al. 4
(VIC). PAINS: Fazenda Amargoso, MG 439 km 16,
29.vii.2004, Melo 1230. PARAOPEBA: 5.x.1957, Heringer 9521 (GH, UB); id., 10.x.1959, Heringer 9505
(UB, US); id., 14.viii.1968, Pereira 10728 (HB, MBM);
id., Fazenda das Pindaíbas, 28.vi.1959, Heringer 7053
(UB, US); id., Fazenda do Rasgão, 20.iv.1954, Heringer 3498 (HB); id., 10.viii.1956, Heringer 5328 (UB,
US); id., Horto Florestal de Paraopeba, 11.v.1974,
Martinelli & Silva 319 (RB). PATOS DE MINAS:
23.viii.1950, Duarte 2762 (BHCB, RB). PERDIZES:
córrego da Aparecida, Unidade de Conservação do
Galheiro, CEMIG, 28.v.1994, Neto & Werneck 1261
(BHCB). SABARÁ: Serra Rachada, 25.viii.2003, Faria
& Versieux 167 (RFA). STA BÁRBARA: Caraça,
22.vii.1972, Emygdio et al. 3579 (R); id., 7.vii.1974,
15
Reitz 7651 (CTES, HBR). SANTANA DO RIACHO:
caminho a Lapinha, 19°10′S 43°41′W, 11.ii.1991, Arbo
et al. 4888 (CTES); id., PN Serra do Cipó, 3 km da
Portaria do Alto Palácio do IBAMA, 25.viii.1992,
Pereira et al. 1053 (BHCB); id., 27.viii.1992, Pereira
et al. 1048 (BHCB); id., rodovia Belo Horizonte–
Conceição do Mato Dentro, 10.ix.1987, Wanderley &
Yano 10707 (SP); id., km 125, 26.iv.1991, Pirani et al.
12310 (SPF); id., Serra do Cipó, km 117, 27.iv.1978,
Martinelli 4297 (RB); Serra do Cabral: c. 2 km N de
Joaquim Felício, 10.iii.1970, Irwin et al. 27369 (GH,
MO, NY, RB, UB, US); id., c. 15 km NW de Joaquim
Felício, 7.vii.1985, Wanderley et al. 817 (SP); Serra do
Cipó, v.1933, Costa 34 (R); id., vii.1949, Vidal s.n. (R
193027); id., vii.1956, Duarte 2703 (RB); id.,
27.viii.1998, Filho et al. s.n. (BHCB 43374); Serra da
Moeda, 5.ix.1968, Pereira 10730 (HB); Serra de Saramenha, viii.1895, Schwacke s.n. (RB 112219); Serra
de Tipahy, 28.viii.1895, Schwacke 11609, 11610 (RB).
SERRO: Milho Verde, 30.iv.2000, Goldschmidt et al.
s.n. (VIC 24843); id., 24.vii.2002, Mota 1846 (BHCB).
SETE LAGOAS: IPEACO, 26.viii.1969, Silva 350 (US).
TIRADENTES: Serra de São José, 11.x.1987, Alves 61
(RB). PARÁ – Cupary river, plateau between the
Xingu and Tapajós rivers, 18.ix.1931, Krukoff 1222
(GH, NY); Estação Ecológica do Jarí; 0°75′S 52°30′W,
17.x.1987, Beck et al. 134 (NY); Itaibuna, estrada
Santarém–Cuiabá, BR 163, km 794, Serra do
Cachimbo, 23.iv.1983, Amaral et al. 927 (NY); Rio
Curuaúna, Cachoeira do Portão, região do Planalto de
Santarém, 30.x.1954, Fróes 31269 (NY, R); Rio Jari,
estrada Monte Dourado–Caracurú, 11.xi.1967,
Oliveira 3594 (NY); Rios Pacaja e Muirapiranga,
2°33′–50′S 50°38′–50′W, 21.ix.1965, Prance et al. 1409
(NY); Rio Xingú, gleba Bacaja, lote 88, below mouth of
rio Bacaja, 3°22′20″S 50°47′50″W, 21.xi.1980, Prance
et al. 26373 (SEL); rodovia Belém–Brasília, km 92,
24.viii.1959, Kuhlmann & Jimbo 110 (SP); Serra do
Cachimbo, iii.1957, Sick s.n. (HB 4604); 25–35 km de
Tucurui, 3°56′S 49°49′W, 4.xi.1981, Daly et al. 1209
(NY). RONDÔNIA – Porto Velho–Cuiabá, Nova Vida,
20.ix.1962, Duarte & Appa 7028 (RB); road Guajará
Mirim to Abunã, km 12, basin of Rio Madeira,
5.viii.1968, Prance et al. 6799 (NY); vicinity of Sta
Bárbara, 15 km east of km 117, 14.viii.1968, Prance &
Ramos 6931 (NY). RORAIMA – estrada Boa Vista–
Venezuela, 5 km S de Rio Surumu, Serra Pacaraima,
2.xii.1977, Steward et al. 194 (NY). SÃO PAULO –
BOTUCATU: 22.xii.1970, Gottsberger 882 (US). CAMPINAS: 20.i.1875, Mosén 3929 (GH foto). CORUMBATAÍ:
Reserva Biológica da UNESP, 15.ii.1996, Paula 1105
(VIC). ITAPETININGA: viii.1901, Wettstein & Schiffner
809 (WU). ITÚ: viii.1967, Friedrich s.n. (HBR
46060). PIRASSUNUNGA: das Emas, 22°2′S 47°30′W,
21.iv.1995, Batalha et al. 401 (SP). SALTO GRANDE: rio
Paranapanema, vii.1901, Wettstein & Schiffner 227
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
16
A. P. G. DE FARIA ET AL.
(WU). SÃO PAULO: 3.ii.1929, Smith & Kuhlmann 1801
(F, GH); id., Butantã, 30.viii.1946, Joly s.n. (SPF
16760); id., Reserva da cidade universitária Armando
Salles de Oliveira, 23°33′S 46°43′W, 3.v.1994, Dislich
80, 81 (SPF); São Paulo, chácara dos Morrinhos, s.d.,
Kruse & Pickel 4629 (SP); São Paulo, Cidade Universitária, jardim do Departamento de Botânica, Instituto de Biociências da USP, 12.vii.1993, Pirani et al.
s.n. (SPF 78037); id., cultivada no Horto Oswaldo
Cruz, 10.ix.1924, Gehrt 13179 (GH, NY, SP); id.,
Jardim Botânico, 18.viii.1980, Wanderley 210 (SP);
id., jardim do Butantã, campus da USP, 1983, Hylio
et al. s.n. (SPF 34337); id., mata do Instituto de Biociências, 15.vii.1994, Batalha & Mello 1 (SPF); id.,
nativa do Jardim Botânico, 31.vii.1939, Handro s.n.
(SP 40195); id., Orchidário, 15.viii.1939, Foster 342
(GH, NY, R); id., Parque Estadual das Fontes do
Ipiranga, Jardim Botânico, 6.vii.1979, Wanderley 116
(SP); id., 16.vii. 2003, Faria & Sousa 148 (RFA); São
Paulo, Pinheiros, vii.1885, Loefgren 2590 (GH foto,
SP); São Paulo, Pirajussara, 3.viii.1930, Gehrt &
Kuhlmann 42 (CTES, SPF); São Paulo, Santo Amaro,
20.vii.1942, Krieger 178 (SP). SOROCABA: 1.viii.1961,
Seidel 29 (HBR). TOCANTINS – MATEIROS: 10°33′S
46°45′W, 8.v.2001, Proença et al. 2510 (MBM, UB).
COLÔMBIA, META – cordilhera La Macarena,
macizo Renjifo, 1950–51, Idrobo & Schultes 925 (GH).
MAGDALENA – PUEBLO BELLO: Sierra Nevada de Sta
Marta, viii.1946, Foster & Smith 1465 (GH). EL
SALVADOR, MORAZAN – rio Negro, 13°59′27″N
88°7′59″W, 24.iii.2002, Monro et al. 3805 (MO).
GUYANA, CUYUNI–MAZARUNI – BARTICA: 12–15
miles from the town, 28.viii.1935, Potter 5332 (GH);
Utshe river area, along trail leading to Venezuela,
5°45′N 61°9′S, 25.v.1990, Mac Dowel & Gopaul
2847 (NY). KAMIKUSA: Mazaruni river, 19.xii.1922,
Leng 385 (NY); Partang river, Merume montains,
23.vi.1960, Maguire & Maguire 45905 (NY). POTARO–
SIPARUNI – Iwokrama Rain Forest Reserve, 4°42′N
58°42′W, 18.ix.1995, Clarke 155 (SEL). UPPER
DEMERARA–BERBICE – Demerara river, ix.1881,
Jenman 4153 (NY); Mabura Hill, 5°20′N 58°40′W,
4.vi.1994, Kellof et al. 1019 (SEL). UPPER TAKUTU–
UPPER ESSEQUIBO – 2°11′N 59°11′W, 19.ix.1993,
Henkel et al. 3069 (SEL). FRENCH GUIANA,
CAYENNE – Montagne de Kaw, 12.xii.1954, Cowan
38751 (NY). SAÜL: vicinity of Eaux Claires, 3°37′N
53°12′W, 3.xi.1992, Mori et al. 22763 (NY); id.,
4.ix.1994, Mori et al. 23781 (NY). HONDURAS,
COMAYAGUA – rio Tepemechin, c. 6 km north of Pito
Solo, 15.iv.1951, Williams & Molina 18008 (GH).
CORTÉS – Ocote Arrancado, 50 km N Lago de Yajoa,
xi.1980, Nelson et al. 5946 (MO). EL PARAÍSO – Ojo
de Água, 27.ii.1947, Standley 4720 (F). OLANCHO –
Poncaya, 4.iii.1982, Blackmore & Heath 1997 (MO).
MÉXICO, QUINTANA ROO – 1 km al leste de Chanca
Veracruz, 7.v.1983, Cabrera & Durán 4646 (MO).
PERU, MADRE DE DIOS – 39 km SW de Puerto
Maldonado, 12°50′S 69°20′W, 9.x.1985, Smith et al.
637 (NY). SURINAME, NICKERIE – área of Kabalebo
Dam Project, 4°–5°N 57°30′–58′W, 17 ix.1980,
Lindman et al. 412 (GH, NY); Lucie Rivier, below
confluence of Oost Rivier, 3°20′N 56°40′W, 6.vii.1963,
Maguire et al. 53996 (NY); 9 km north of Lucie Rivier,
12 km west Oost Rivier, 3°36′N 56°30′N, 18.vii.1963,
Maguire et al. 54258 (NY); id., 21 vii.1963, Maguire
et al. 54333 (NY); 12 km north of Lucie Rivier, 3 km
south of Juliana top, 3°36′–3°41′N 56°30′–56°34′W,
29.viii.1963, Irwin et al. 55153 (MO, NY); Tafelberg
(Table Mountain), 10.viii.1944, Maguire 24276 (NY);
Tanjimama River, 20.xi.1954, Mennega M487 (NY).
TRINIDAD, ARIMA – iv.1874, O. Kuntze 988 (NY);
Arima Valley, Sta Isabella trace, 24.i.1959, Aitken s.n.
(TRIN 23688). CARAPICHAIMA – 23.iii.1892, Alexander 5723 (GH, NY). SAINT ANDREW – Balandra,
March 1922, Freeman s.n. (GH foto), Long Stretch,
13.vii.1976, Adams 14011 (NY, TRIN). SANDRE
GRAND – SANGRE GRANDE: Tractor trace, Melajo
forest, 2.viii.1955, Martinez s.n. (TRIN 23648); Valencia, near Quare Reservoir, 11.v.1933, Broadway s.n.
(GH). VENEZUELA, AMAZONAS – ATABAPO: between
Duida and Marahuaca, near base of Duida, 3°34′N
65°32′W, 28.x.1988, Liesner 25569 (MO); id., Cerro
Marahuaca, 3°43′N 65°31′W, 28.iii.1985, Steyermark
& Holst 130896 (MO). ATURES: Rio Coro-Coro, west of
Serrania de Yutage, 5°38′N 66°7′30″W, 19.ii.1987,
Holst & Liesner 3096 (MO). RÍO NEGRO: along the Rio
Mawarinuma, Neblina Massif, c. 7 km east-north-east
of Puerto Chimo, 0°50–51′N 66°2–6′W, vii.1984,
Davidse & Miller 27185 (MO); id., c. 1 km north-east
of San Carlos de Rio Negro, 20 km south of confluence
of Rio Negro and Brazo Casiquire, 1°56′N 67°3′W,
7.v.1979, Liesner 7266 (MO); id., cerro de La Neblina,
camp IV, 15 km north-north-east of Pico Phelps,
0°51′N 65°57′W, iii.1984, Liesner 16813 (MO, NY).
ARAGUA – COLONIA TOVAR: 1854–55, Fendler 1352
(GH); id., north-east of Maya, 7 km by air south-east
of Colonia Tovar, 10°21′30″N 67°14′30″W, 24.iii.1980,
Steyermark & Liesner 121854 (MO). APURE – PÁEZ:
selva Cutufí, between Rio Cutufí and Rio Sanare,
7°9–11′N 71°56–58′N, xi.1982, Davidse & González
21955 (MO). BOLÍVAR – Cerro Guaiquinima: 5°45′N
63°35′W, 26.v.1978, Steyermark et al. 117471 (MO);
Cerro Socopa, between estado Falcón and Lara,
10°29′N 70°48′W, 29.vi.1979, Liesner et al. 8350 (MO).
GRAN SABANA: c. 10 km south-west of Karaurin
Tepui, 5°19′N 61°3′W, 22.iv.1988, Liesner 23597 (MO);
Miamo, Hato de Nuria, 25.i.1961, Steyermark 88852
(NY); Ptari-tepuí, xi.1944, Steyermark s.n. (GH);
represa Guri, 7°46′N 63°W, 31.iii.1981, Liesner &
González 11036 (MO); Rio Caura, 2–8 km S del Salto
Para (Las Pavas), 6°13′N 64°28′W, 10.v.1982 Morillo
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
AECHMEA SUBGENUS MACROCHORDION
& Liesner 9107 (MO); Rio Paragua, Salto de Auraima,
10.iv.1943, Killip 37339 (GH); Serra Imataca, Rio
Toro, N of El Palmar,12.xii.1960, Steyermark 87984
(NY). ROSCIO: alrededores del Río Apongueo,
17.xi.1981, Burandt et al. V11120 b (MO); id., ‘El
Abismo’, Rio Samay, 4°23′N 64°38′W, 26.x.1985, Holst
et al. 2501 (MO). DELTA AMACURO – TUCUPITA:
5–14 km east-south-east of Los Castillos de Guayama,
8°28′N 62°17′W, 1979, Davidse & González 16421
(MO). MIRANDA – Cerro Sipapo (Paráque), 2.ii.1949,
Maguire & Politi 28786 (NY); morros de al Guairita,
16.viii.1975, Berry 1025 (NY). PORTUGUESA – 30 km
west of Guanare by air, along Rio Tucupido, 9°2′N
70°1′W, 11.iii.1982, Liesner et al. 12488 (MO). SUCRE
– peninsula de Araya, 20 km north-west of Cariaco by
air, 10°38′N 63°40′W, v.1981, Liesner & González
12050 (MO). TÁCHIRA – 10 km east of La Fundacíon,
around Represa Dorada, iii.1981, Liesner & González
10169 (MO); Serra El Casadero, 13 km north of
Rubio, between La Dantas and Las Adjuntas, 7°43′N
72°23′W, 12.xi.1979, Steyermark et al. 120133 (MO).
ZULIA – COLÓN: intersection of Rio Catatumbo and
the La Fria Maracaibo highway, 9°7′N 72°37′W,
29.vi.1980, Davidse et al. 18818 (MO, SEL).
Notes: With one of the widest geographical distributions within Bromelioideae, A. bromeliifolia var.
bromeliifolia is the best represented taxon of the
complex in herbarium collections. This taxon also
presents an extensive list of synonyms. Here, we
kept the 14 taxonomic synonyms considered by
Smith & Downs (1979) and propose two more:
A. bromeliifolia var. angustispica (see previous discussion) and A. lagenaria (a synonym of A. lamarchei prior to this study). This last is justified
because A. lagenaria shows many typical characters
of A. bromeliifolia (e.g. leaf spines > 3 mm and truncate floral bracts) that are not founded in A. lamarchei. Despite the great morphological variation of
some vegetative structures (e.g. rosette and leaf
shape), A. bromeliifolia var. bromeliifolia is delimited by a set of reproductive characters, including
flowers 1.4–1.6 cm long, densely lanate, depressed
ovate floral bracts, with truncate or truncate emarginate apices and yellow petals with fimbriate
ligulae at the distal end of the lateral folds (Fig. 5B,
C, E). This species also resembles A. triangularis in
having leaf spines ⱖ 3 cm, emarginate, symmetric to
slightly asymmetric sepals, connate to the middle,
and spatulate petals with emarginated apices
(Fig. 5D, E).
2.2. Aechmea bromeliifolia var. albobracteata Philcox,
Ashingtonia 1(8): 92. 1974 (Fig. 10E, F)
Type: Brazil, Mato Grosso, 1 km E do km 264, estrada
Xavantina–Cachimbo, 12°49′S, 51°46′W, 21.iii.1968,
Philcox & Ferreira 4605 (Holotype K! photo).
17
Description: LEAF SHEATHS with adaxial surface vinaceous or purpureous, abaxial surface green. LEAF
BLADES concolorous green, margins serrate; spines
black, 3–6 mm long. INFLORESCENCE with peduncle
green; peduncle bracts white. FLORAL BRACTS green.
SEPALS green. PETALS yellow greenish.
Distribution and habitat: Aechmea bromeliifolia var.
albobracteata occurs from Argentina and Paraguay
to central, south and south-eastern Brazil, in Mato
Grosso, Mato Grosso do Sul, Minas Gerais, São Paulo
and Paraná states (Fig. 6). It grows at 100–900 m
altitude, in semi-deciduous forest, cerrados and
campos rupestres.
Conservation status: Least Concern (IUCN, 2001).
Specimens examined: ARGENTINA, CORRIENTES –
ITUZAINGÓ: Isla Apipé Grande, Puerto Mora,
1.xii.1973, Krapovickas et al. 24279 (CTES). SANTO
TOMÉ: estancia Garrunchos, ayo Chimiray, 6.ii.1972,
Krapovickas et al. 21051 (CTES). MISIONES –
POSADAS: 28.xi.1907, Ekman 1211 (GH photo). SAN
IGNÁCIO: Teyucuaré, 12.xi.1976, Quarín 3502 (CTES).
BRAZIL, MATO GROSSO DO SUL – CORUMBÁ:
estrada para o Jacadigo, 24.x.1992, Wendt et al. 261
(COR). MINAS GERAIS – BARROSO: mata do
Baú, 26.viii.2001, Assis & Ladeira 180, 195 (CESJ);
id., 20.x.2001, Forzza et al. 1931 (CESJ). LAMBARI:
6.viii.1967, Pereira 10615 (HB, MBM). SANTO
ANTÔNIO DO ITAMBÉ: próximo ao Parque Estadual do
Pico do Itambé, 29.viii.2003, Faria & Versieux 171
(RFA). SÃO GONÇALO DO RIO PRETO: P.E. do Rio
Preto, 18°6′8″S 43°20′22″W, 14.vi.2002, Lombardi
et al. 4857 (BHCB). UBERLÂNDIA: clube caça de pesca
Itororó, 30.viii.2001, Araújo s.n. (HUFU 33309).
VIÇOSA: in cultivation on Horto Botânico da UFV,
18.ix.1979, Filho s.n. (VIC 6337); id., 20.ix.1999,
Goldschmidt s.n. (VIC 23790). PARANÁ – GUAÍRA:
Sete Quedas, 27.i.1962, Reitz & Klein 12151 (HBR).
GUARATUBA: 30.vii.1972, Hatschbach 29834 (MBM).
JAGUARIAÍVA: 26.x.1910, Dusén 10779 (GH photo); id.,
17.viii.1914, Dusén 15446 (GH photo); id., 18.xi.1914,
Dusén 16072 (GH, MO, NY); id., 27.viii.1915, Dusén
s.n. (GH photo); id., Parque Estadual do Cerrado,
17.x.2000, Lisingen 57 (MBM). PIRAÍ: 3.viii.1973,
Seidel 658 (HB). PONTA GROSSA: Parque Velha,
2.x.1965, Hatschbasch 12876 (MBM). SENGÉS:
Fazenda Morungava, rio do Funil, 10.ix.1959, Hatschbasch 6318 (HBR, MBM); Serrinha, 22.x.1908, Dusén
7024 (GH photo, NY). VENTANIA: 15.vi.1960, Seidel
s.n. (HBR 46065); Vila Velha, 19.xii.1903, Dusén
2799 (R); id., 27.viii.1939, Foster 409 (GH, R); id.,
28.viii.1939, Kuhlmann s.n. (SP 41547). SÃO PAULO –
ANHEMBI: Fazenda Barreiro Rico, 6.x.1956, Kuhlmann 3998 (SP). BROTAS: NW of the intersection of
the road Brotas–Itirapina, 22°17′S 47°56′W, 16.vi.
1961, Eiten et al. 2956 (SP). ITAPURA: 29.ix.1940,
Foster 1099 (GH). MOJI–GUAÇÚ: Reserva Florestal
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
18
A. P. G. DE FARIA ET AL.
perto de Pádua Salles, 19.vii.1955, Handro 506 (SP).
SÃO CARLOS: parque da Universidade Federal de
São Carlos, viii.2000, Moura s.n. (RB 376129). SÃO
PAULO: nativa do Jardim Botânico, 8.viii.1932,
Hoehne s.n. (SP 29789). PARAGUAY, ALTO PARANÁ –
estancia Bertoni (Rio Paraná), 25°38′S 54°36′W,
22.vii.1994, Zardini & Florentín 40049 (MO).
AMAMBAY – CABALLERO: 20.i.1889, Morong 523 (NY).
CAAGUAZÚ – GUAYAQUÍ: between Coronel Oviedo and
Caaguazú, 25°29′S 56°11′W, 26.viii.1993, Zardini &
Tilleria 37032 (SEL). CAAZAPÁ – TAVAÍ: 26°10′S
55°27′W, 28.x.1988, Zardini 7680 (CTES, MO); id.,
29.x.1988, Zardini 7741 (CTES, MO). CORDILLERA –
Cerro Zanja Jhú, 1 km east of road from route 1 to
Atyra, 25°13′S 57°9′W, 25.vi.1988, Zardini 5150 (MO);
eastern side of Rio Piribebuy basin, 17 km west of
Arroyos y Esteros, 25°8′S 57°15′W, 23.xii.1989,
Zardini & Velázquez 17163 (MO); road Eusébio Ayala,
PN Vapor Cué, Arroyo Yakarey, 25°30′S 56°50′W,
7.vii.1990, Zardini & Velázquez 21916 (MO). ITACURUBI DE LA CORDILLERA: 3.ii.1991 Till 6015 (WU). EL
CHACO – riacho Negro, 14.ix.1893, Lindman A2155
(GH photo). GUAIRÁ – VILLARICA: 9.xi.1928, Jorgensen 3961 (F, GH photo, NY); Cosme, Villarica–
Caaguazu, iii.1876, Balansa 706 (MO photo, NY
photo); Rio Chololo, Cordillera de Peribébuy,
15.ix.1883, Balansa 4748 (MO photo, NY photo).
ITAPÚA – PIRAPÓ: 6.viii.1893, Lindman A 1893 (GH
photo). PARAGUARÍ – Arroyo Moopicua, 26°20′32″S
57°8′45″W, 28.vii.1994, Zardini & Guerrero 40176
(SEL); Tebicuarymí river, 1 km north of Villa Florida,
26°23′S 57°8′W, 25.v.1993, Zardini & Guerrero 35829
(WU). SAN PEDRO – Alto Paraguay, Primavera,
Tajoma Isla, 22.viii.1959, Woolston 1372 (NY); Yaguareté forest (Sustainable Forest Systems site),
23°49′46″S 56°12′48″W, 24.viii.1995, Zardini &
Vargas 43519 (MO); id., 23°47′56″S 53°13′6″W,
24.viii.1995, Zardini & Vargas 43674 (MO); id.,
1.viii.1996, Zardini & Guerrero 45463 (SEL).
Notes: In the taxonomic treatment for Aechmea subgenus Macrochordion, Smith & Downs (1979) disregarded the relevance of white peduncle bracts for the
recognition of A. bromeliifolia var. albobracteata, and
this type specimen was included in the list of examined material for A. bromeliifolia var. bromeliifolia.
Almost all herbarium material listed as A. bromeliifolia var. albobracteata in this study was previously
identified as var. bromeliifolia. In revising specimen
identifications, besides the colour of the peduncle
bracts, we also considered information about peduncle
and floral bract coloration and the geographical origin
of the specimen.
3. Aechmea lamarchei Mez, in Martius, Eichler &
Urbain, Fl. Bras. 3(3): 370. 1892. (Figs 7A–F, 10G, H)
Type: Brazil, without specific locality, Lamarche
Hortus in Morren Hortus s.n., x.1877 (Holotype: LG!
photo).
= Macrochordion lamarchei (Mez) L.B.Sm. &
W.J.Kress, Phytologia 66(1): 77. 1989. H.E.Luther &
E.Sieff, Selbyana 15 (1):65. 1994, pro syn.
= Aechmea maculata L.B.Sm., Smithsonian Misc.
Collect. 126:15, 227, f. 107. 1955. Type: Brazil, Minas
Gerais, Belo Horizonte, Pico da Piedade, 10.vii.1940,
Foster 561 (Holotype: GH!, Isotype: US! photo,
BHCB!), syn. nov.
= Macrochordion maculata (L.B.Sm.) L.B.Sm. &
W.J.Kress, Phytologia 66(1): 77. 1989, syn. nov.
= Aechmea chlorophylla L.B.Sm., Smithsonian
Misc. Collect. 126:14, 227, f. 108. 1955. Type: Brazil,
Espírito Santo, Sta Teresa, 6.viii.1940, Foster 830
(Holotype: GH!, Isotype: US! photo), syn. nov.
= Macrochordion chlorophylla (L.B.Sm.) L.B.Sm. &
W.J.Kress, Phytologia 66(1): 77. 1989, syn. nov.
Description: HERB epiphytic, rupicolous or terrestrial,
30–80.5 cm high. ROSETTE funnelform or wide-funnelform, with 15–30 leaves. LEAF SHEATHS 10–20 ¥ 2.5–
10.5 cm, elliptic to narrow–elliptic, adaxial surface
vinaceous, abaxial surface green or sometimes
reddish; LEAF BLADES 17–112.5 ¥ 2–5.5 cm, linear,
concolorous green or sometimes concolorous reddish,
apex acute or obtuse, apiculate, margins serrate or
densely serrate; spines castaneous, 1–3 mm long.
INFLORESCENCE with spike 3.5–12.5 ¥ 1.5–4 cm;
peduncle green or reddish, 27–73.5 cm long, white
floccose; peduncle bracts imbricate or subdense, divergent toward the apex of the peduncle, entire or denticulate toward the apex, dark pink or red, 6.5–
14 ¥ 2–4 cm, apex acute or acuminate. FLOWERS 2.2–
2.8 cm long. FLORAL BRACTS 1–2.3 ¥ 1.5–2 cm, ovate
or widely ovate, shorter to about equaling the sepals,
coriaceous with thin apex, reddish or castaneous,
white floccose or apressed lepidote, apex obtuse or
emarginate, apiculate. SEPALS 8–14 ¥ 4–8 mm, distinctly asymmetric, connate near the base or sometimes to the middle, yellow, yellow–greenish or
reddish, covered with white appressed scales, apex
obtuse, muticous or minutely apiculate. PETALS
17–23 ¥ 5–6 mm, lingulate, yellow, apex obtuse, erect
or slightly divergent; fimbriate ligulae about halfway
along the lateral folds. STAMENS with filaments
9–16 mm long; anthers 5–9 mm long. STIGMA c. 2 mm
long. OVARY c. 5 mm long, bearing a distinct epigynous tube; ovules c. 0.5 mm long. FRUITS greenish;
seeds c. 4 mm long, pale brown.
Distribution and habitat: Aechmea lamarchei is distributed in Minas Gerais and inland areas of Espírito
Santo state (Fig. 4B). It grows at 500–1700 m altitude, in dense ombrophile forest, semi-deciduous
forest and campos rupestres.
Conservation status: Least Concern (IUCN, 2001).
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
AECHMEA SUBGENUS MACROCHORDION
19
Figure 7. Aechmea lamarchei A, habit. B, detail of leaf margin. C, flower with floral bract. D, floral bract. E, sepals. F,
petal and stamens. A, B, E, F (Faria et al. 161). C, D (Irwin et al. 29141).
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
20
A. P. G. DE FARIA ET AL.
Specimens examined: BRAZIL, ESPÍRITO SANTO –
CACHOEIRO DE ITAPEMIRIM: Vargem Alta, Morro de
Sal, 2.ix.1948, Brade 19414 (RB); id., 16.viii.1981,
Ferreira 1836 (RB). CAPARAÓ: Fazenda Laranja da
Terra, 11.viii.1972, Heringer 12163 (HB). CARIACICA:
31.i.1986, Seidel 1024 (RB). DOMINGOS MARTINS:
9.vii.1939, Foster 176 (GH, R); id., in cultivation,
REBIO de Sapitanduva, 2.viii.1992, Hatschbach
57141 (MBM); id., without date, G. Hatschbach
58052 (MBM); id., próximo à cidade de Campinho,
27.viii.1974, Martinelli 415 (RB); id., 12.ix.1975,
Martinelli 771 (RB); id., rod. BR 262, próximo ao Rio
Araguaia, 12.x.1992, Hatschbach et al. 57996 (MBM).
ITARANA: Jatiboca, Fazenda Stuhr, 7.viii.2002, Kollmann et al. 5671 (MBML). MARECHAL FLORIANO:
Fazenda do Sr. Bullbach, próximo a Todos os Santos,
9.ii.1975, Martinelli & Gurken 560 (RB); id., Sítio de
Almir Bresson, 27.vi.1994, Gomes 2022 (VIES). STA
LEOPOLDINA: Timbuí Seco, 8.vii.1988, Krause s.n.
(MBML 5013). STA MARIA DE JETIBÁ: São José do
Rio Claro, propriedade Luzineide Butke, 5.viii.2001,
Santos et al. 19 (MBML). STA TERESA: in cultivation,
Museu de Biologia Mello Leitão, 28.ix.1998,
Fernandes 2571, 2572, 2573, 2574 (MBML); Sta
Teresa, divisa com município de Sta Leopoldina, cabeceira do rio Novo, 7.xi.1986, Martinelli et al. 11886
(RB); Sta Teresa, Estação Biológica da Caixa D’água,
19.v.1998, Fernandes 2510 (MBML); id., in cultivation, UFRJ, 2.ix.2005, Wendt et al. 385 (RFA); Sta
Teresa, Estação Biológica de Sta Lúcia, 20.viii.1985,
Boone 701 (NY, MBML); id., 25.viii.1986, Martinelli
et al. 11617 (RB); id., 31.viii.1998, Varassin 78
(MBML, RFA); id., 7.xii.2000, Wendt et al. 390, 391
(RFA); id., 26 ix.2002, Faria et al. 49, 50 (RFA);
id., 13.viii.2003, Faria et al. 161 (RFA); Sta Teresa,
Fazenda Tabajara: 28.xi.1985, Piziolo 249 (CEPEC,
NY, MBML); Sta Teresa, mata ao redor da pousada
Paradiso, 28.ix.2002, Faria et al. 70 (RFA); Sta Teresa,
pátio do Museu de Biologia Mello Leitão, 27.viii.1985,
Fernandes 1441 (MBML); id., 27.viii.1986, Fernandes
1439 (MBML); Sta Teresa, Pedra da Onça, propriedade de Antônio Rocon, 13.vi.2000, Demuner et al.
1103 (MBML); Sta Teresa, Penha, 21.viii.1984, Boone
304 (MBML), id., 8.viii.1988, Loss &. Santos s.n.
(MBML 5043); Sta Teresa, São João de Petrópolis,
Colégio Agrotécnico, 19°49′S 40°43′W, 16.vii.1968,
Gottsberger 18-16768 (SP); Sta Teresa, Valsugana
Velha, 16.ix.1985, Boone 769 (NY, MBML); id.,
5.ix.2001, Kollman & Bausen 4531 (MBML); id.,
14.ix.2001, Kollman & Bausen 4565 (MBML). MINAS
GERAIS – ABRE CAMPO: 1° Distrito, Fazenda Cachoeira, 20°14′16″S 42°29′31″W, 21.xii.2000, Pereira
29/59 (RFA). CAETÉ: Serra da Piedade, 26.viii.1987,
without collector (BHCB); id., 1.vi.2001, Mota &
Marques 325 (BHCB). CARANGOLA: Fazenda da
Grama, c. 5 km north of ‘Mato Virgem’, 4.ii.1930,
Mexia 4316a (GH, US). CARATINGA: Fazenda Montes
Claros, 19.iv.1984, Andrade & Lopes 148 (BHCB).
CATAS ALTAS: Serra do Caraça, 15.iii.1998, Vasconcelos s.n. (BHCB 41412). CONCEIÇÃO DO MATO DENTRO:
Serra do Cipó, north of Belo Horizonte, vii.1940,
Foster 630, 637 (GH, US). CORONEL FABRICIANO:
Mata do Limoeiro, 15.x.1950, Magalhães 5838 (US).
CORONEL PACHECO: 6.viii.1945, Heringer 1968 (SP).
DESCOBERTO: Reserva Biológica da Represa do
Grama, 24.vi.2000, Salimena et al. 196 (CESJ); id.,
19.vii.2001, Castro 531 (CESJ); id., 23.iii.2002, Forzza
et al. 2114 (CESJ). DIONÍSIO: Fazenda do José Gomes,
19°49′44″S 42°40′02″W, 8.vi.2000, Temponi et al. 121
(VIC). FERVEDOURO: Parque Estadual da Serra do
Brigadeiro, 15.iv.1993, Leme et al. 2172 (SEL); id.,
13.iv.1995, Paula 1032 (VIC), id., 21.vi.1996, Paula
1130 (VIC); Fervedouro, Reserva Municipal do
Córrego da Água Limpa, 42°21′S 20°46′W, 6.viii.1989,
Leoni 832 (RB); id., 18.viii.1992, Nahoum s.n. (SEL
076176). ILHÉU: Fazenda da Tabunha, 18.viii. 1930,
Mexia 4972 (F, GH, MO, US, VIC). JEQUERI: área de
inundação da Usina de Providência, 19.xi.1997,
Salino 3763 (BHCB). LEOPOLDINA: 25.xi.1978, Seidel
783 (HB). MARLIÉRIA: Parque Estadual Rio Doce,
2.vii.1993, Borba & Costa s.n. (BHCB 30674). OURO
BRANCO: Serra do Ouro Branco, 20.xi.2002, Paula s.n.
(VIC 27382). RIO POMBA: estrada Rio Pomba a Barbacena, 21.vii.1969, Braga 60 (RB). RIO VERMELHO:
trilha Alto da Serra do Ambrósio, c. 8 km de Coluna,
18°9′92″S 42°58′15″W, 1.viii. 2000, Costa & Fiaschi
387 (SP). SANTANA DO RIACHO: Serra do Cipó,
próximo km 127, 15.vii.1977, Martinelli 2631 (RB,
UB). STA BÁRBARA: Brumal, Fazenda Bocaina,
23.v.1997, Vasconcelos s.n. (BHCB 42717); Sta
Bárbara, Serra do Caraça, 16.iv.1933, Mello Barreto
2113 (BHCB); id., 7.vii.1974, Reitz 7650 (HBR);
id., 24.v.1987, Paula s.n. (BHCB 9739); id., 2.x.1998,
Mota et al. s.n. (BHCB 43448); id., caminho para
a gruta do Padre Caio, 23.v.1987, Zappi & Scatena
10972 (SPF); Sta Bárbara, Parque Nacional do
Caraça, caminho para a cascatona, 24.iv.1990,
Leme et al. 1546 (RB); Sta Bárbara, Serra do Caraça,
caminho para a Capelinha, SW de Catas Altas,
20°5′S 43°27′W, 18.ii.1991, Arbo et al. 5309 (CTES,
SPF, US); Sta Bárbara, Serra do Caraça, próximo ao
colégio do Caraça, 19.vii.1977, Martinelli 2687 (RB,
US); Serra do Espinhaço, base of Serra do Caraça,
26.i.1971, Irwin et al. 29141 (UB); Serra do Espinhaço, c. 17 km of Serro, on road to Diamantina,
27.ii.1968, Irwin et al. 20994 (UB); Vale do Rio
Piranga, estrada entre Porto Firme e Guaraciaba,
12.x.1996, Palhais & Paula 44 (VIC). VIÇOSA: Agricultural College lands, 24.vi.1930, Mexia 4789-a (GH);
id., in cultivation on Horto Botânico da UFV,
29.vi.1979, Vidal et al. 482 (VIC); id., 13.vi.1985,
Vidal & Vidal s.n. (VIC 9386); id., 15.vii.1995, Paula
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AECHMEA SUBGENUS MACROCHORDION
1043 (VIC); id., 5.vii.1999, Paula & Goldschmidt s.n.
(VIC 23647).
Notes: In the original descriptions of A. chlorophylla
and A. maculata, Smith (1955) proposed an affinity
of both species with A. bromeliifolia. Multivariate
analyses, however, showed that A. bromeliifolia is
distinguishable and that there is a close relationship
between A. chlorophylla, A. maculata and A. lamarchei. The difficulties to distinguishing these three
taxa morphologically were already noted in the work
of Smith & Downs (1979). Most of the diagnostic
characters presented for A. lamarchei in the key to
species of subgenus Macrochordion can also be
observed in A. chlorophylla and/or in A. maculata
(e.g. leaf blade shape, inflorescence indument type,
floral bract apex, length of fusion among sepals). We
also ascertained that the measures (length and wide)
of leaves, leaf blade spines, inflorescences, sepals
and petals of A. chlorophylla herbarium specimens
examined fit into those described for A. lamarchei.
Leaves and peduncle bracts spotted with red, a diagnostic character cited for A. maculata in the Smith
& Downs (1979) key, is not clearly defined in the
morphological description of this species, which mentions dense and coarse purple spots only on the
internal face of the sheaths. Purple sheaths,
however, are commonly observed in the subgenus.
Other characters employed by Smith & Downs
(1979) to distinguish A. lamarchei from A. chlorophylla and A. maculata have been shown to be variable and of dubious interpretation. The white lanate
inflorescence cited for A. lamarchei was not observed
in herbarium material examined in this study.
Instead, we observed floccose to apressed lepidote
inflorescences and these characters are cited in the
key to identify A. maculata and A. chlorophylla,
respectively. Leaf blades linear vs. ligulate, used in
the key to differentiate A. lamarchei from A. chlorophylla and A. maculata is not adequate (the three
species show linear blades). The distinction between
lacerate (cited for A. chlorohylla) and fimbriate petal
appendages (cited for A. lamarchei and A. maculata)
were not detected in our study and all scales were
classified as fimbriate. The connation of the appendages to the petals (slightly above the base cited for
A. lamarchei, at the base for A. chlorophylla and
above the base, near the middle of the claw for
A. maculata) also leads to misinterpretation. We
observed that, in all these species, the fimbriate
scales are inserted well above the base of the petals
and approximately halfway along the lateral folds.
Another character employed by Smith & Downs
(1979) to distinguish these taxa was the length of
fusion among the sepals (half connate for A. lamarchei, approximately half connate for A. maculata
and connate at base for A. chlorophylla). Examina-
21
tion of herbarium material, however, showed considerable variation of this character within A. lamarchei. Considering that these species have been hard
to distinguish from each other historically, and the
close relationships suggested among them by the
multivariate analysis, we propose A. chlorophylla
and A. maculata as synonyms of A. lamarchei. We
consider the coriaceous, ovate to widely ovate floral
bracts with obtuse or emarginate thin apices and the
large flowers > 2 cm (Fig. 7C, D) relevant to identify
A. lamarchei (these characters are also present in A.
chlorophylla and A. maculata) and to distinguish
this species from other taxa of the subgenus.
4. Aechmea maasii Gouda & W.Till, Bromelia 4(1): 4.
1997 (Figs 8A–F, 10I, J)
Type: Brazil, Espírito Santo, Linhares, Reserva Florestal Vale do Rio Doce (CVRD), 1992 (bloomed in
cultivation at University of Utrecht Botanic Gardens,
xi. 1995), Maas s.n. (Holotype: U not seen, Isotypes:
CVRD!, WU! photo).
Description: HERB epiphytic or terrestrial, 30.5–75 cm
high. ROSETTE funnelform, wide-funnelform or tubulose with 20–25 leaves. LEAF SHEATHS 7.5–21 ¥ 4.5–
12.5 cm, orbicular to transversely elliptic, adaxial
surface vinaceous, abaxial surface green. LEAF
BLADES 22.5–44.5 ¥ 2–5 cm, linear, concolorous green
or sometimes yellowish, apex obtuse or acute, apiculate, margins serrate or densely serrate; spines castaneous, 1–3 mm long. INFLORESCENCE with spike
3.5–9 ¥ 1.5–3.5 cm; peduncle green or reddish, 25.5–
70 cm long, white floccose; peduncle bracts imbricate
or subdense, erect or divergent toward the apex of the
peduncle, margins entire or denticulate toward the
apex, red, 6–15 ¥ 1.5–3 cm, apex acuminate. FLOWERS
1.8–2.1 cm long. FLORAL BRACTS 8–12 ¥ 13–16 mm,
depressed ovate, shorter than the sepals, coriaceous,
castaneous or green with apex and margins reddish,
densely white floccose, apex truncate, apiculate.
SEPALS 9–11 ¥ 5–7 mm, distinctly asymmetric,
connate near the base, yellow, yellow–greenish or
reddish, covered with white appressed scales, apex
obtuse, muticous or minutely apiculate. PETALS
15–17 ¥ 4–5 mm, lingulate, yellow, apex obtuse, erect
or slightly divergent; fimbriate ligulae about halfway
along the lateral folds. STAMENS with filaments
9–13 mm long; anthers 4.5–5 mm long. STIGMA
1–2 mm long. OVARY 3–4 mm long, bearing a distinct
epigynous tube, ovules 0.5–1 mm long. FRUITS
globose, red; seeds c. 4 mm long, pale brown.
Distribution and habitat: Aechmea maasii occurs in
coastal areas of Espírito Santo and Rio de Janeiro
states (Fig. 4C). It grows at 5–100 m altitude, in herbaceous, shrubby and wooded restingas and florestas
de tabuleiros.
Conservation status: Vulnerable (IUCN, 2001).
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
22
A. P. G. DE FARIA ET AL.
Figure 8. Aechmea maasii. A, habit. B, detail of leaf margin. C, flower with floral bract. D, floral bract. E, sepals. F, petal
and stamens. A, B (Faria 177). C–F (Faria 147).
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
AECHMEA SUBGENUS MACROCHORDION
23
Figure 9. Aechmea triangularis. A, habit. B, flower with floral bract. C, floral bract. D, sepals. E, petal and stamens.
(Wendt et al. 392).
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A. P. G. DE FARIA ET AL.
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
AECHMEA SUBGENUS MACROCHORDION
25
Figure 10. Aechmea subgenus Macrochordion taxa. A, B, Aechmea alba (Faria et al. 139); C, D, Aechmea bromeliifolia
var. bromeliifolia (Faria & Sousa 148): inflorescences and flowers. E, F, Aechmea bromeliifolia var. albobracteata (Faria
& Versieux 171); G, H, Aechmea lamarchei (Faria et al. 161, 212, respectively): habit and detailed inflorescences. I, J,
Aechmea maasii (Faria 177); K, L, Aechmea triangularis (Wendt et al. 392): detailed inflorescences and flowers. All
photographs by the authors, except A. bromeliifolia var. albobracteata by L. M. Versieux.
䉳
Specimens examined: BRAZIL, ESPÍRITO SANTO –
ALFREDO CHAVES: 18.vii. 1990, Marbach 09 (VIES).
ARACRUZ: Comboios, 7.i.1992, Pereira et al. 2528
(VIES); id., 8.i.1992, Pereira et al. 2570 (VIES); id.,
28.vii.1992, Pereira et al. 3629 (VIES); Aracruz,
Retiro, 29.iv.1992, Pereira et al. 3352 (VIES); id.,
6.v.1992, Pereira et al. 3398 (VIES); Aracruz, Vila do
Riacho, área 106 da Aracruz Celulose S.A., 19.ii.1992,
Pereira et al. 2733 (VIES). CONCEIÇÃO DA BARRA:
área 126 da Aracruz Celulose S.A., 26.ii.1992, Pereira
et al. 2874 (VIES); id., área 213 da Aracruz Celulose
S.A., 24.iii.1992, Pereira et al. 3037 (VIES); id.,
25.iii.1992, Pereira et al. 3069 (VIES); id., área 157 da
Aracruz Celulose S.A., 27.iii.1992, Pereira et al. 3263
(VIES); id., área 100 da Aracruz Celulose S.A.,
25.viii.1992, Pereira et al. 3758 (VIES); Conceição da
Barra, Itaúnas, 20.v.1999, Hatschbach et al. 69179
(MBM). GUARAPARI: Parque Estadual Paulo César
Vinha, 25.iii. 1991, Gomes 1482 (VIES); id., 9.vii.1991,
Gomes 1678 (VIES); id., 22.i.1998, Gomes 2399
(VIES); id., 28.vi.1998, Gomes 2431 (VIES); Guarapari, praia do Morro, 30.viii.1974, Martinelli &
Gurken 479 (RB); Guarapari, restinga de Setiba,
26.vii.1990, Pereira 2146 (VIES). ITAPEMIRIM:
10.viii.1980, Gurken 6 (HB); id., cultivada por L. C.
Gurken, 6.vii.2001, Faria et al. 41 (RFA); Itapemirim,
in cultivation, UFRJ, Faria 147 (RFA); Itapemirim,
estrada Marataízes–Piúma, c. de 4 km ao norte da
ponte sobre o Rio Itapemirim, 20°58′16″S 40°48′42″W,
21.vi.1996, Mello-Silva et al. 1191 (SPF). LINHARES:
Reserva Biológica de Comboios, 21.iii.1991, Gomes
1470, 1473 (VIES); Linhares, Reserva Florestal Cia.
Vale do Rio Doce (CVRD), 30.i.1972, Sucre 8287 (RB);
id., 1.ii.1972, Sucre 8395 (RB); id., 17.i.1975, Peixoto
& Peixoto 392 (RB); id., 10.v.1977, Martinelli et al.
1901 (RB); id., 27.ix.1978, Martinelli 5007 (RB); id.,
17.xii.1981, Lima 1714 (RB); id., 20.iv.1983, Farney
et al. 287 (RB); id., 15.ix.1987, Martinelli 12207 (RB);
id., 17.ix.1987, Martinelli 12221 (RB); id., 18.ix.1987,
Martinelli 12237 (RB); id., 22.ii.2000, Folli 3576
(CVRD); id. 29.vi.2000, Folli 3643 (CVRD); id.,
11.ix.2001, Folli 4046 (CVRD); id., 16.x.2001, Folli
4095 (CVRD, SEL); id., 17.xii.2001, Folli 4153
(CVRD); id., 18.ix.2003, Folli 4606 (CVRD); id.,
30.vii.2004, Folli 4887 (CVRD); Linhares, Reserva
Florestal de Linhares, 23.ii.1999, Folli 3355 (CVRD);
id., 24.ii.1999, Folli 3357 (CVRD); id., 27.ii.1999, Folli
3500 (CVRD); id., 9.xii.1999, Folli 3528 (CVRD); id.,
31.v.2000, Folli 3593 (CVRD); Linhares, Reserva Florestal de Sooretama, 9.viii.1965, Belém 1518 (UB); id.,
11.vii.1969, Sucre 5452 (RB); id., 15.v.1977, Martinelli
et al. 2075 (RB); id., 12.v.1985, Martinelli et al. 10989
(RB); Linhares, Regência, 19.ix.1991, Gomes 1610
(VIES). SÃO MATEUS: 4.x.1964, Seidel 529 (HBR); id.,
ligação BR 101 a Ponta do Ipiranga, 14.x.1992,
Hatschbach et al. 58056 (MBM); id., 10.xi.1993,
Hatschbach & Silva 60071 (CEPEC, MBM). SERRA:
Bicanga, 6.v.1993, Pereira 4551 (VIES); id., 27.v.1993,
Pereira et al. 4563 (VIES). RIO DE JANEIRO – BÚZIOS:
x.2002, Wendt et al. 443 (RFA); id., in cultivation,
UFRJ, 5.ix.2003, Faria 177, 178 (RFA). CABO FRIO:
17.vii.1967, Pereira 10607 (HB); id., in cultivation,
Marie Selby Botanical Gardens, 30.xii.1997, Berg s.n.
(SEL 078423); Cabo Frio, Morro do Gavião, 13.x.1968,
Sucre 3918 (RB); Cabo Frio, south-west of Armação de
Búzios, 3.ii.1995, Till et al. 11131 (WU). CASIMIRO DE
ABREU: distrito de Barra de São João, 1953, SegadasVianna et al. 277 (R); id., 5.ix.1953, Segadas-Vianna
et al. 967-I (GH, R, RB); id., 3.vii.1963, Andrade &
Emmerich 1590 (R); id., 14.viii.1986, Costa et al. 7
(RB). MACAÉ: restinga de Macaé, 30.ix.1974, Martinelli & Gurken 506 (RB). SAQUAREMA: 12.ix.1986,
Farney & Caruso 1183 (R, RB); id., 9.viii.2003, Faria
et al. 149 (RFA); id., Ipitangas, 8.ix.1987, G. Martinelli et al. 12185 (RB).
Notes: In the original description, Gouda & Till
(1997) proposed an affinity of A. maasii with A. bromeliifolia and A. triangularis. Phenetic analyses,
however, indicate that A. maasii is closely related to
A. alba and A. lamarchei (Figs 1, 2). Aechmea
maasii shows a great ecological plasticity and phenotypic variation in vegetative structures (Scarano
et al., 2002, 2009), but can be characterized by the
flowers with yellow corollas and depressed ovate,
truncate–apiculate and densely floccose floral bracts
(Figs 8D, 10I, J). The species is also characterized
by its restricted distribution in areas of Espírito
Santo and Rio de Janeiro states (Fig. 4C). Aechmea
maasii resembles A. alba and A. lamarchei in
having leaf spines ⱕ 3 mm long, obtuse, distinctly
asymmetric sepals, lingulate petals with obtuse
apices and fimbriate ligulae approximately halfway
along the lateral folds (Fig. 8B, E, F).
5. Aechmea triangularis L.B.Sm., Smithsonian Misc.
Collect. 126: 19, 224, f. 106. 1955 (Figs 9A–E, 10K, L)
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 1–27
26
A. P. G. DE FARIA ET AL.
Type: Brazil, Espírito Santo, Sta Teresa, 7.viii.1940
(bloomed in cultivation 22.iv.1941), Foster 829 (Holotype: GH!, Isotype: GH!).
= Macrochordion triangularis (L.B.Sm.) L.B.Sm. &
W.J.Kress, Phytologia 66(1): 77. 1989. H.E.Luther &
E.Sieff, Selbyana 15 (1):65. 1994, pro syn.
= Aechmea kautskyana E. Pereira & L.B.Sm.,
Bradea 3: 46–7, 49. 1980. Type: Brazil, Espírito
Santo, Conceição do Castelo, Forno Grande,
8.viii.1979, Kautsky 643 (Holotype HB!, Isotype US!
photo), syn. nov.
= Macrochordion kaustyana (E. Pereira & L.B.Sm.)
L.B.Sm. & W.J.Kress, Phytologia 66(1): 77. 1989, syn.
nov.
Description: HERB epiphytic, 23.5–50 cm high.
ROSETTE funnelform or wide-funnelform with 15–20
leaves. LEAF SHEATHS 8–18 ¥ 4.5–20 cm, orbicular to
elliptic, adaxial surface purple, abaxial surface green.
LEAF BLADES 10–72 ¥ 1.5–6 cm, linear or linear triangular, concolorous green, sometimes purple spotted,
apex caudate, recurved, margins serrate or sparsely
serrate; spines black, 3–7 mm long. INFLORESCENCE
with spike 3–6 ¥ 2–3 cm; peduncle vinaceous,
20–41 cm long, white floccose; peduncle bracts subdense and divergent, margins denticulate toward
the apex, dark pink or red, 4–7.5 ¥ 1.3–3 cm, apex
acute or acuminate. FLOWERS 1.6–2 cm long. FLORAL
BRACTS 8–11 ¥ 11–13 mm, depressed ovate, shorter
than the sepals, coriaceous, red to the middle and
castaneous toward the apex, white floccose, apex
truncate, apiculate. SEPALS 6–8 ¥ 4 mm, symmetric or
slightly asymmetric, connate to the middle, vinaceous
or castaneous with red margins, covered with white
appressed scales, apex emarginate, minutely apiculate. PETALS 12–16 ¥ 4 mm, spatulate, dark blue with
white base, apex emarginate, erect; fimbriate ligulae
about halfway along the lateral folds. STAMENS with
filaments c. 1 cm long; anthers 4–5 mm long. STIGMA
c. 1 mm long. OVARY c. 4 mm long, bearing a short
epigynous tube, ovules c. 0.5 mm long. FRUITS and
seeds not observed.
Distribution and habitat: Aechmea triangularis is
endemic to Espírito Santo state (Fig. 4D) and is
poorly represented in herbarium collections. It grows
at 650–1000 m altitude, in dense ombrophile forest.
Conservation status: Endangered (IUCN, 2001).
Specimens examined: BRAZIL, ESPÍRITO SANTO –
NOVA LOMBARDIA: divisa com mata do IBDF,
31.x.1984, Hoffmann 228 (MBML). STA LEOPOLDINA:
cabeceira do Rio Bonito, divisa com município de Sta
Teresa, 12.xii.1988, Fernandes 2284 (MBML). STA
TERESA: Estação Biológica de Sta Lúcia, 18.xii.1998,
Varassin et al. 59 (MBML); id., 23.iii.1999 Varassin
et al. 66 (MBML); id., 7.xii.2000, Wendt et al. 392
(RFA). VARGEM ALTA: 21.xii.1976, Pereira s.n. (HB
65953).
Notes: Aechmea triangularis is the most distinctive
taxon in Aechmea subgenus Macrochordion and can
be easily recognized by both vegetative (e.g. leaves
with caudate and recurvate apices) and reproductive
(e.g. dark blue corolla) characters (Figs 9A, 10K, L).
As noted by Pereira (1980) on the original description,
A. kautskyana also has blue petals. The author,
however, stressed the presence of dimorphic leaf
blades as a relevant character in considering A.
kautskyana distinct from A. triangularis. Differences
between inner and outer leaf blades were also
observed in some herbarium and living specimens of
A. triangularis. This characteristic, however, is not
present in young plants as occurs in true dimorphic
leaves. In mature plants of A. triangularis, the inner
leaves of the rosette are narrower and suddenly
become broader, and the same seems to occur in
A. kaustskyana. Until this study, A. kaustkyana
remained known only from the type specimen. The
close morphological similarity indicated in the
phenetic analyses (Figs 1, 2) supports the synonymy
proposed for these species.
ACKNOWLEDGEMENTS
We thank the Brazilian Forestry Service (IBAMA)
and the Brazilian Research Council (CNPq) for the
field collection permits; the numerous colleagues who
assisted us with plant collection; H. Luther, who
provided some of the plant material; L.O.F. de
Sousa and L.M. Versieux for miscellaneous help; G.
Gonçalves for the drawings; the US National Science
Foundation (DEB-0129446) for funding; the Brazilian
Council for Graduate Studies (CAPES) for a research
grant to A. P. G. de Faria; and CNPq for a productivity grant to T. Wendt. This paper is part of a PhD
thesis undertaken at the Postgraduate Program in
Botany of the Universidade Federal do Rio de Janeiro
by the first author.
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